Proceedings of the California Academy of Sciences, 4th series

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

| APR 11 2002

|

Volume 52
2000-2001

California Academy of Sciences
Golden Gate Park
San Francisco, California

SCIENTIFIC PUBLICATIONS DEPARTMENT

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Katie Martin, Managing editor
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ISSN 0068-547X

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16.

TABLE OF CONTENTS
VOLUME 52

MUNIAIN, CLAUDIA and ANGEL VALDES. Rostanga byga Er. Marcus, 1958 from Argentina:
Redescription and comparison to Rostanga pulchra MacFarland, 1905 (Mollusca,

Mudibranchia: Dortdina). (ilus.):-February 23,2000) of) 02 6 COL 2s See ea 1-10
KOCIOLEK, J. PATRICK. Valve ultrastructure of some Eunotiaceae (Bacillariophyceae), with
comments on the evolution of the raphe system. (Illus.) February 23, 2000.......... 11-21
HOARE, RICHARD D. Silurian polyplacophora and rostroconchia (Mollusca) from northern
amionnia: .(Ullus, pReonlary 25, 2000. 2.82.06 oo aceas oka ease ee eee ee ees eed 23-31
ALMEDA, FRANK. New Costa Rican and Panamanian species of Miconia (Melastomataceae:
Micomecte)r sllinus:\: Wied: 2000) » cere cee et ett go Sie Peas uo don aS ae ee ee 33-54

DREWES, ROBERT C. and JEAN-LUC PERRET. A new species of giant, montane
Phrynobatrachus (Anura: Ranidae) from the central mountains of Kenya. (IIlus.)

Pa pce rae aes eA a Al cua el eg sia eS ee nana conc (Pie. de x RNG, ¥ vio Wes Sages ee 55—64
WILLIAMS, GARY C. Two new genera of soft corals (Anthozoa: Alcyoniidae) from South
Africa, with a discussion of diversity and endemism in the Southern African octocorallian
AMURTE MULLIS Syed UL ye, DONO case che) eh eye ae a EN one iiaxtt aves 2 -/RE:2) seta ee tone 65-75
VALDES, ANGEL and ORSO ANGULO CAMPILLO. Redescription and reassessment of Cadlina
luarna (Ev. Marcus and Er. Marcus, 1967), comb. nov. (Mollusca, Opisthobranchia,

“VT ISTaliey) Pe (CITI Pe inae2 oYe7 0) 0) OI pe a a ne rk ee Pe Ey ier 77-85
WILLIAMS, GARY C. and JOHN STARMER. A new genus and species of soft coral (Octocorallia:
Micyonudac) trom South Africa. (illus.) October 18, 2000) 2... bo ne ee eee 87—96
ALMEDA, FRANK. The hexandrous species of Topobea (Melastomataceae). (Illus.)

Oehe pe 1S CUD 2 U6 eee oe Be Oe eS ee ee ee erm ree yn 97-109

. GOSLINER, TERRENCE M. and DAVID W. BEHRENS. Two new species of Chromodorididae

(Mollusca: Nudibranchia) from the tropical Indo-Pacific, with a redescription of Hypselodoris
doljusi (Provot-r ol, 1933): @llus.) October 18; 2000. ...722.. 6 ea oe eee bs ee 111-124

. NG, HEOK HEE and CARL J. FERRARIS, JR. A review of the genus Hemibagrus in southern

Asia, with descriptions of two new species. (Illus.) December 7, 2000............. 125-142

. DANIEL, THOMAS F., KEVIN BALKWILL, and MANDY-JANE BALKWILL. Chromosome numbers

alsouth Atrican Acanthaceae. (illus.) December 7, 2000...... 2 22 cence ie ws 143—1358

. WILLIAMS, GARY C. A new species of the soft coral genus Eleutherobia Pitter, 1900

(Coelenterata: Alcyoniidae) from the Tonga Islands. (Illus.) December 7, 2000...... 159-169

. ELWOoD, HILLARY R., ANGEL VALDES, and TERRENCE M. GOSLINER. Two new species of

Aldisa Bergh, 1878 (Mollusca, Nudibranchia) from the tropical Indo-Pacific. (Illus.)

Pee rearan ic tea) ese he ice hy eet, eco tee cra heen yaad at's obthshin wha,-e: Sya> kee Sw a 171-18)
BEHRENS, DAVID W. and ANGEL VALDES. The identity of Doris (s.l.) species MacFarland,
1966 (Mollusca, Nudibranchia, Discodorididae): A persistent mystery from California solved.
CEIIDS J JE ATOVE TSTE 22100) I gee gett ett in oy Par Ri err 183-193
WILLIAMS, GARY C. and SUSAN A. LITTLE. A new species of the soft coral genus
Eleutherobia Pitter, 1900, (Octocorallia: Alcyoniidae) from South Africa. (Illus.)

JEOTTTISE OLAV NOL ae) Sh See ern eran, ae ne ag Ree eee ee ae arene ee 195—208

. WILLIAMS, GARY C. First record of a bioluminescent soft coral: Description of a disjunct

population of Eleutherobia grayi (Thomson and Dean, 1921) from the Solomon Islands, with
a review of bioluminescence in the Octocorallia. (Illus.) August 21, 2001......... 209-225

. LEE, WELTON L. Four new species of Forcepia (Porifera, Demospongiae, Poecilosclerida,

Coelosphaeridae) from California, and synonymy of Wilsa de Laubenfels, 1930, with
toreepia. Carter, 1874. (lus.)'October 26, 2000... 0. a5. ogee Sele Pew eee aes 227-244

. DE FREITAS, SERGIO and NORMAN D. PENNY. The green lacewings (Neuroptera: Chrysopidae)

ol Bravalian agro-ccosystems. (lllus.) October 26, 2001... 2... 2.005 ceed. ee ween. 245-397

PROCEEDINGS TABLE OF CONTENTS, VOL. 52

20. SLOWINSKI, JOSEPH B., SAMRAAT S. PAWAR, HTUN WIN, THIN THIN, SAI WANNA GYI, SAN
LWIN Oo, and HLA TUN. A new Lycodon (Serpentes: Colubridae) from Northeast India and
Myanmar (Burma). (lhis:))October 265 2001S 2 Ere asso. ae ee 397-405

21. IWAMOTO, TOMIO and KEN J. GRAHAM. Grenadiers (Families Bathygadidae and Macrouridae,
Gadiformes, Pisces) of New South Wales, Australia. (Illus.) November 13, 2001 ....407—S09

22. ALMEDA, FRANK. The octandrous and dodecandrous species of Topobea (Melastomataceae)
in Mexico-and Central Amenca, (llus;)November 13,2001). es 94... eee 511-548

InGdexeto W/OlUMeS2 Aer Ae ores carte = oe A ee ee a 549-558

PROCEEDINGS, VOLUME 52

ERRATA

Page (Para.) Line Corrections
Number 18
233: (12) 3 “CASIZ 53465” to read “SIO-BIC P1368”
236 (1) ] “CASIZ 35961” to read “SBMNH 345543”
Number 19
280 (3) 9 “walkeriana’ to read “walkerina”
282 (1) l “walkeriana’ to read “walkerina”
283 (3) 3 “walkeriana’ to read “walkerina”
283 (8) 4 “walkeriana’ to read “walkerina”

6 “walkeriana’ to read “walkerina”
310 (1) 1 “vittatus”’ to read “vittata”
eh (3}) 9 “vittatus” to read “vittata”
310 (4) “vittatus” to read “vittata”
310 (4) l “atus” to read “ata”

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 1, pp. 1-10. 6 figs., 1 table. February 23, 2000

Rostanga byga Er. Marcus, 1958 from Argentina: Redescription
and Comparison to Rostanga pulchra MacFarland, 1905
(Mollusca, Nudibranchia, Doridina)

by LUUU
Claudia Muniain

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia, ” Avda. Angel Gallardo 470 (1405),
Buenos Aires, Argentina

and

Angel Valdés
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Rostanga byga Marcus, 1958 is redescribed based on the examination of the type material
and newly collected specimens from Golfo San José, Chubut, Argentina. The external col-
oration, radular morphology and reproductive system differ significantly from specimens
of Rostanga pulchra MacFarland, 1905 collected from Monterey Bay, California (type lo-
cality), and examined for comparison. This latter species was probably recorded from Ar-
gentina (Camarones Bay, Chubut) as well. The geographic range of R. byga is extended
from northern Brazil to Patagonia. Rostanga pulchra appears to have a disjunct geo-
graphic range in North and South America.

RESUMEN

Se redescribe la especie Rostanga byga Marcus, 1958 a partir del estudio del material tipo y
de varios especimenes recolectados por primera vez en el Golfo San José, Chubut, Argen-
tina. La coloracion del cuerpo, morfologia radular y sistema reproductivo difieren
significativamente de especimenes de Rostanga pulchra MacFarland, 1905 recolectadosen
la localidad tipo, la Bahia de Monterey, California y examinados para su comparacion.
Probablemente, esta ultima especie ha sido también citada para Argentina (Bahia
Camarones, Chubut). La distribucion de RX. byga se extiende desde el norte de Brasil hasta
la Patagonia. Rostanga pulchra parece tener dos areas de distribucion disjuntas, en Norte y
Sur América.

The genus Rostanga Bergh. 1879 comprises species of caryophyllidia-bearing dorids character-
ized by having short rhinophores with few lamellae. inner radular teeth folded inwards and outer teeth
elongate and denticulate.

So far, two species of Rostanga have been reported from South America. Ernest Marcus (1958)
described Rostanga byga based ona single. preserved specimen collected from the intertidal zone of
Ilhabela. Sao Sebastido Island. Brazil. Later, Eveline Marcus (1970) extended the geographical range
of this species to northern Brazil. Rudman and Avern (1989) revised the genus Rostanga from the
Indo-West Pacific. and discussed other species of this genus described from all over the world. These
authors recognized R. byga as a valid species. even though there was very little available information
about it.

MUNIAIN AND VALDES: ROSTANGA BYGA ER. MARCUS, 1958 FROM ARGENTINA 2

Ernest Marcus (1959) reported Rostanga pulchra MacFarland, 1905 from Chile, the first record
of this species (originally described from California), from South America. Years later, Marcus and
Marcus (1969) extended the geographic range of this species to Argentina, based ona single specimen
collected at 102 m depth from Camarones Bay (Chubut). More recent records of R. pulchra from
Chile (Schrédl 1996, 1997) confirmed the presence of this species in the southern Hemisphere. Ac-
cording to the published information, R. pulchra is known to have a disjunct geographic range in
North and South America, with a large gap between the populations.

The present paper includes a complete anatomical study of several specimens of Rostanga col-
lected from Argentina, and a comparison with material of R. pulchra from the type locality. In addi-
tion, the type material of R. byga is re-examined and compared to the original description.

The material examined is deposited at the Museo Argentino de Ciencias Naturales “Bernardino
Rivadavia” (MACN), the Department of Invertebrate Zoology and Geology of the California Acad-
emy of Sciences (CASIZ), and the American Museum of Natural History (AMNH).

SPECIES DESCRIPTIONS

Family Rostangidae Pruvot-Fol, 1951
Genus Rostanga Bergh, 1879

Rostanga pulchra MacFarland, 1905
Figs. 1,2; 3A

Rostanga pulchra MacFarland, 1905:40-41; 1906:119—122, pl. 24, fig. 8, pl. 18, figs. 18-21, pl. 21, fig. 109;
1966:165—169, pl. 25, fig. 7, pl. 29, figs. 7-10, pl. 35, figs. 1-16; Er. Marcus, 1959:35—37, 106, figs. 65—68;
Er. Marcus, 1961:15, pl. 3, figs. 46-49; Marcus and Marcus, 1969:20-21; Marcus and Marcus,
1970:202—203; Thompson, 1975:489; Rudman and Avern, 1989:330; Schrédl, 1996:22; Schrédl,
1997:38-42.

MATERIAL EXAMINED. — Monterey Bay, California, 17 August 1978, three specimens 11, 12
and 16 mm preserved length, collected by G. McDonald (CASIZ 069163); 24 August 1978, three
specimens 14, 15 and 18 mm preserved length, collected by G. McDonald (CASIZ 070734).

EXTERNAL MORPHOLOGY. — The body shape is oval, somewhat elevated. The dorsum is cov-
ered with cup-shaped caryophyllidia, about 150 um long. The rhinophoral and branchial sheaths are
also surrounded by caryophyllidia. Each caryophyllidium consists of an extremely small, rounded,
ciliated tubercle with small marginal cilia and 4-5, taller, thin spicules surrounding the tubercle. The
living animals (Fig. 1) vary from orange-yellow to bright red in color. Some specimens may have
small, brown spots scattered on the dorsum. The mantle margin is surrounded by small, white spots.
The perfoliate rhinophores have the same color as the dorsum and are composed of 9 vertical, trans-
verse lamellae. The gill has the same color as the rest of the body, and is composed of 6—10 bipinnate
branchial leaves. The anal papilla lies within the circlet of the branchial plume.

Ventrally, the anterior border of the foot is bilabiate and notched. The foot is wide relative to the
mantle margin. The oral tentacles are well formed, and appear conical in shape.

INTERNAL ANATOMY. — The radular formula is 50 < 48.0.48 ina 16 mm preserved length speci-
men (CASIZ 069163) and 53 x 51.0.51 in an 18 mm preserved length specimen (CASIZ 070734).
The innermost lateral teeth are the shortest of the row. They are thin, folded inwards, and have 6-8
denticles on the inner side of the short cusp (Fig. 2A). The lateral teeth change gradually in size from
the inner to mid-lateral. The mid-lateral teeth are wide, having a long, pointed, primary cusp and a
large secondary cusp situated near the base (Fig. 2B). The outer lateral teeth are very thin and elon-
gate, with a fine brush of 9—10 denticles at the end (Fig. 2C, D). The jaws are solid, with few rodlets of
varying length (Fig. 2E).

3 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. |

FiGurE 1. Rostanga pulchra MacFarland, 1905, living animals from Monterey Bay. showing color variation and an egg
mass. Photograph by Robert Ames (CASIZ photo collection).

In the reproductive system the ampulla is wide and short. It narrows before branching into the
oviduct. which enters the female gland. and the large prostate (Fig. 3A). The prostate is flat and mas-
sive, and has two well-differentiated portions. The prostate narrows and forms the proximal end of the
deferent duct. The distal end of the deferent duct is very long. The penis has no armature. The deferent
duct and vaginal duct meet at a common atrium. The vaginal duct is very long and narrows before
opening into the large. oval bursa copulatrix. At some distance from the vaginal duct insertion, an-
other long duct leads from the bursa copulatrix and connects to the seminal receptacle and the uterine
duct. The seminal receptacle is small and oval. with a slightly pointed end opposite its short stalk.

GEOGRAPHIC RANGE. — This species is known from Alaska (Lee and Foster 1985) to Mexico
(Marcus and Marcus 1970) in the Northern Hemisphere. and from Chile (Ernest Marcus 1959:
Schrédl 1996. 1997) to Argentina (Marcus and Marcus 1969) in the Southern Hemisphere.

Rostanga byga Er. Marcus, 1958
Figs. 3B, 4-6

Rostanga byga Er. Marcus, 1958:22-25, figs. 34-36; Ev. Marcus, 1970:943; Rudman and Avem, 1989:329.
Rostanga cf. pulchra (MacFarland): Muniain, 1997:21

MATERIAL EXAMINED. — HOLOTYPE: IIhabela. Sao Sebastiao Island. Brazil. June 1956. one
specimen 1] mm preserved length, collected under a stone in the intertidal zone (AMNH 3507). Golfo
San José. Chubut, Argentina. November, 1991. one specimen 12 mm preserved length (CASIZ
118014) and one specimen 14 mm preserved length (MACN 34174) collected by C. Muniain; April
1998, one specimen 20 mm long. collected by C. Muniain.

MUNIAIN AND VALDES: ROSTANGA BYGA ER. MARCUS, 1958 FROM ARGENTINA 4

FIGURE 2. Rostanga pulchra MacFarland, 1905, scanning electron micrographs (CASIZ 070734). A. Inner lateral teeth,
scale = 25 um; B. Mid-lateral teeth, scale = 30 um; C. Outer lateral teeth, scale = 60 um; D. Detail of the outermost teeth
denticulation, scale = 7.5 um; E. Jaw, scale = 60 um.

EXTERNAL MORPHOLOGY. — The body shape is oval and somewhat elevated (Figs. +A. 5). The
surface of the mantle is densely covered with caryophyllidia about 100 um long. The rhinophoral and
branchial sheaths are also surrounded by caryophyllidia. Each caryophyllidium consists of a large.
rounded ciliated tubercle. with small marginal cilia and 6 spicules surrounding the tubercle. The
ground color of the mantle is orange in the living animals. lightly spotted with white dots in the middle

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 1

5

fg if

FIGURE 3. Reproductive systems. A. Rostanga pulchra MacFarland, 1905 (C ASIZ 070734). B. Rostanga hyga Er. Marcus,
1958 (CASIZ 118014). Abbreviations: a, ampulla: be, bursa copulatrix: dd, deferent duct: fg, female gland: pr, prostate: s, sem-
inal receptacle: v, vagina.

of the dorsum and between tubercles. The perfoliate rhinophores are pale orange with translucent tips.
and are composed of 14—16 vertical. transverse lamellae (Fig. 4B). The gill has the same color as the
dorsum. and is composed of 10 short, simply pinnate, branchial leaves. The anal papilla lies within the
circlet of the branchial plume. Ventrally. the anterior border of the foot is notched and grooved (Fig.
4C). The foot is completely orange. and narrow relative to the mantle margin. The oral tentacles are
digitiform.

INTERNAL ANATOMY. — The radular formula is 45 = 53.0.53 ina 12 mm preserved length speci-
men (CASIZ 118014) and 58 « 55.0.55 ina 14 mm preserved length specimen (MACN 34174). The
innermost lateral teeth are the shortest of the row. They have a rounded. long cusp with 7—10 short
denticles on the inner edge (Fig. 6A). The lateral teeth change gradually in size from the inner to
mid-lateral. The mid-lateral teeth have a broad triangular base and a pointed primary cusp (Fig. 6B).
These teeth lack a secondary cusp. but have a small lateral wing near the base. The outermost lateral
teeth are very thinand elongate. having a fine brush of 6—10 denticles at the end (Fig. 6C. D). The jaws
are very small. having few irregular rodlets of varying shape and length (Fig. 6E).

In the reproductive system the ampulla is thin and long. It narrows before branching into the ovi-
duct. which enters the female gland. and the prostate (Fig. 3B). The prostate is flat and very large. It
has two well-differentiated portions. In one of them the deferent duct leaves as a narrow long duct. The
penis has no armature. The deferent duct and vaginal duct meet at acommonatrium. The vaginal duct
opens into the large. oval. bursa coopulatrix. Next to the opening of the vaginal duct leads another
duct, which connects to the seminal receptacle and the uterine duct. The seminal receptacle is oval and
conspicuous.

GEOGRAPHIC RANGE. — This species is known from the southwest Atlantic, from the North of
Brazil to Argentina (Chubut).

DISCUSSION
The Argentinean specimens of Rostanga examined in this paper clearly belong to R. byga. The

holotype of R. byga was dissected when it was originally examined, and the radula and reproductive
system were missing. However. there is enough information in the original description (Emest

MUNIAIN AND VALDES: ROSTANGA BYGA ER. MARCUS, 1958 FROM ARGENTINA 6

FIGURE 4. Rostanga byga Er. Marcus, 1958 (MACN 34174). A. Living animal, scale = 3 mm; B. Lateral and overview of the
rhinophores, scale = 2.5 mm; C. Ventral view of the anterior border of foot, scale = 4.5 mm.

Marcus 1958). to compare our specimens withR. byga. According to Ernest Marcus (1958). 2. bygais
a bright, brick red species with dorsal white spots. Our specimens from Argentina have a similar col-
oration. withvery distinctive white dots on the dorsum. Anatomically. in X. bvga the innermost lateral
teeth have a long cusp. longer than the base. and the lateral teeth lack a secondary cusp. Also. in the re-
productive system drawn by Ernest Marcus (1958) for Rk. byga. the ducts leading from the bursa
copulatrix are next to each other, the prostate is large and the ampulla is elongated. All of these charac-
teristics. which are diagnostic of R. byga, are present in our specimens.

Based on the examination of several specimens of R. pu/chra from California and RX. byga from
Argentina. it is clear that they constitute two distinct species (see Table 1). Externally. &. byga is anor-
ange species with white spots on the dorsum. whereas R. pu/chra varies from orange to bright red.
sometimes having black or brown dorsal spots. Also, R. byga has twice as many lamellae in the
rhinophores as R. pulchra. Inaddition, the caryophyllidiaof R. pu/chra have a small ciliated tubercle,
whereas it is very large in R. byga. Internally. the innermost lateral teeth of R. pu/chra have a short

7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52. No. |

cusp. with long denticles.
whereas in RX. byga the cusp of
the innermost teeth is longer.
with shorter denticles. The
mid-lateral teeth of R. pulchra
have a large. secondary cusp
situated under the main cusp.
whereas this secondary cusp is
very small in #. byga. The jaws
of R. pulchra have several
elements regularly arranged.

whereas in RX. byvga the jaws are
very reduced. with only a few. FIGURE 5. Rostanga byga Er. Marcus, 1958, holotype (AMNH 3507).

irregular elements. The ampul-

la of R. pulchra is comparatively shorter and wider than that of R. byga. and the prostate of R. pulchra
appears to be smaller. In addition. the two ducts emerging from the bursa copulatrix of R. pulchra are
separated. whereas they are next to each other in R. byga.

The other Atlantic species of Rostanga is Rostangarubra (Risso. 1818) from the northeast Atlan-
tic. This species is clearly distinguishable from R. pu/chra and R. rubra by the presence of very wide
inner lateral teeth, with strong denticles, and elongate outermost teeth with a single denticle (Thomp-
sonand Brown 1984). This is very different from bothR. pulchra and R. byga that have narrower inner
teeth and outer teeth with numerous denticles.

The present record of R. byga constitutes the first account of this species from the temperate wa-
ters of the Argentinean biogeographic Province, near the limit with the Magellanic Province. Records
of the genus Rostanga from the Magellanic Province have beenassigned to R. pu/chra (Ermest Marcus
1959: Marcus and Marcus 1969: Schrodl 1996, 1997). The first of them (Ernest Marcus 1959), in-
cludes a complete anatomical description of the specimens collected from Chile. According to this de-
scription. there is no question that the specimens studied belong to R. pu/chra. The morphology of the
radula. with a short cusp on the innermost lateral teeth and a large secondary cusp on the mid-lateral
teeth is characteristic of this species. The other records from Chile (Marcus and Marcus 1969: Schrédl
1996. 1997). include brief descriptions without anatomical information. and it is difficult to determine
the identity of the animals studied. The record from Argentina (Marcus and Marcus 1969) is particu-
larly problematic. not only because it is based ona single preserved specimen. but also because it was
collected from a locality very close to the limit between the Magellanic and Argentinean Provinces.
According to Marcus and Marcus (1969), the specimen was collected at 102 m depth, and had a very
large radular formula (85 =< 90.0.90). larger than that of either R. pulchra or R. byga. The animal was
preserved. so there was no information available on the external coloration. However. Marcus and
Marcus (1969) indicated that there were several dark spots on the dorsum of the preserved animal.
whichis characteristic of R. pu/chra. With the available information we can not determine the identity
of this specimen. but it is very likely that it belongs to R. pulchra.

The record of Rostanga byga from Northern Chubut is very close to the southern boundary of the
Argentinean biogeographic Province with the cold waters of the Magellanic Province. According to
Carcelles and Williamson (1951). this boundary is situated in Golfo Nuevo, about 42°30'S. However.
it is not unusual to find warm-water species of opisthobranch mollusks in this transitional area. Other
temperate or warm-water species with a similar distributional pattern to Rk. bvga have been recently
studied from Argentina (see Muniain 1997; Muniain and Ortea 1998: Muniain and Ortea, in press).
On the other hand. Carcelles and Williamson (1951) and Muniain (1997). showed that the Magellanic

r

MUNIAIN AND VALDES: ROSTANGA BYGA ER. MARCUS, 1958 FROM ARGENTINA

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9 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 1

FIGURE 6. Rostanga byga Er. Marcus, 1958, scanning electron micrographs (CASIZ 118014). A. Inner lateral teeth,
scale = 30 um; B. Mid-lateral teeth, scale = 43 um; C. Outer lateral teeth, scale = 75 um: D. Detail of the outermost teeth
denticulation, scale = 10 um; E. Jaw, scale = 15 um.

species collected from Northern Chubut to Rio de la Plata (Argentinean Province) always occur in
deeper and colder waters (100-200 m). This phenomenon is probably related to a northern extension
of the Falklands Current in deeper waters (Carcelles and Williamson 1951: Muniain 1997). The fact
that the specimen of R. pulchra reported by Marcus and Marcus (1969) from this area was collected
from deep (102 m), and therefore colder waters, seems to support this hypothesis.

MUNIAIN AND VALDES: ROSTANGA BYGA ER. MARCUS, 1958 FROM ARGENTINA 10

ACKNOWLEDGMENTS

The authors would like to recognize the support of several individuals. Terry Gosliner allowed us
to study material of the California Academy of Sciences and reviewed the manuscript. James
Cordeiro and Paula Mikkelsen (American Museum of Natural History). provided us with the type ma-
terial of Rostanga byga. which was photographed at the Photography Department of the Museo
Argentino de Ciencias Naturales. Dong Lin, Photography Department of the California Academy of
Sciences, printed the scanning electron micrographs.

This paper has been supported in part by the Fundacion Antorchas and Conicet. through a re-
search subsidy and a postdoctoral fellowship to the first author, the Ministerio de Educaciony Cultura
of Spain (SEUI). through a postdoctoral fellowship to the second author, and the California Academy
of Sciences.

LITERATURE CITED

CARCELLES, A. AND S. WILLIAMSON. 1951. Catalogo de los moluscos marinos de la Provincia Magallanica.
Revista del Museo Argentino de Ciencias Naturales, Zoologia 2:225-383.

LEE, R. AND N. Foster. 1985. A distributional list with range extensions of the opisthobranch gastropods of
Alaska. Veliger 27:440-448.

MacFARLAND, F. M. 1905. A preliminary account of the Dorididae of Monterey Bay, California. Proceedings of
the Biological Society of Washington 18:35—34.

. 1906. Opisthobranchiate Mollusca from Monterey Bay, California, and vicinity. Bulletin of the United

States Bureau of Fisheries 25:109-151, pls 18-31.

. 1966. Studies of opisthobranch mollusks of the Pacific Coast of North America. Memortrs of the Califor-
nia Academy of Sciences 6:1—546, pls 1-72.

Marcus, ERNEST. 1958. On Western Atlantic Opisthobranchiate gastropods. American Museum Novitates
1906: 1-80.

. 1959. Lamerallariaceaund Opisthobranchia. Report of the Lund University Chile Expedition 1948-49,

N° 36. Lunds Universitets Arsskrift (Ny. Foljd), Adv. 2, 55:1-133.

. 1961. Opisthobranch mollusks from California. Veliger 3:1—85, pls. 1-10.

Marcus, EvVELINE. 1970. Opisthobranchs from Northern Brazil. Bulletin of Marine Science 20:922-951.

Marcus, EVELINE AND ERNEST MARcus. 1969. Opisthobranchian and lamellarian gastropods collected by
“Vema.” American Museum Novitates 2368: 1-31.

. 1970. Some gastropods from Madagascar and West Mexico. Malacologia 10:181—223.

MunlaIn, C. 1997. Moluscos Opistobranquiosde Argentina: revision taxonomicay relacion de ecologia quimica
en algunas especies patagonicas. Unpublished Ph. D. Thesis, Universidad de Oviedo, Spain.

MUunliAIn, C. AND J. ORTEA. 1998. The taxonomic status and redescription of Polycera marplatensis Franceschi,
1928 (Nudibranchia: Polyceratidae) from Argentina. Veliger 41: 142-147.

.In press. New records of the genus Berghia Trinchese, 1877 (Opisthobranchia: Aeolidiudae) from Ar-
gentina, with description of a new species. Avicennia.

RUDMAN, W. AND J. AVERN. 1989. The genus Rostanga Bergh, 1879 (Nudibranchia: Dorididae)in the Indo-West
Pacific. Zoological Journal of the Linnean Society 96:281-338.

SCHRODL, M. 1996. Nudibranchia y Sacoglossa de Chile: morfologia externa y distribucion. Gayana Zoologica
60:17-62.

. 1997. Range extensions of Magellanic nudibranchs (Opisthobranchia) into Peruvian Faunal Province.
Veliger 40:38-42.

THompsON, T. E. 1975. Doridnudibranchs from eastern Australia (Gastropoda, Opisthobranchia). Journal of Zo-
ology 176:477-517.

THompson, T. E. AND G. H. Brown. 1984. Biology of opisthobranch molluscs. Volume II. The Ray Society,
London. 229 pp.

© CALIFORNIA ACADEMY OF SCIENCES. 2000
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 2, pp. 11-21, 35 figs. February 23, 2000

Valve Ultrastructure of Some Eunotiaceae (Bacillariophyceae),
with Comments on the Evolution of the Raphe System

by
J. Patrick Kociolek a
Diatom Collection, California Academy of SciencdlAR O57
Golden Gate Park, San Francisco, California 94118 yee

Ultrastructural observations on valve features of five diatom species from the family
Eunotiaceae are presented, including four from the Amazon basin and one from Califor-
nia. Of the Amazonian taxa, Peronia brasiliensis Hustedt has a straight raphe system posi-
tioned on the valve face and a central sternum, Eunotia synedraeformis Hustedt and E.
curvula Hustedt have raphe systems predominantly towards the valve mantle, as well as a
central sternum. Eunotia conversla Hustedt has a raphe system positioned more on the
valve face and a central sternum. Specimens of Amphicampa eruca Ehrenberg from Cali-
fornia have a very short raphe on the mantle and lack a central sternum. These data, in ad-
dition to previously published observations, point to the diversity of raphe types in the
Eunotiaceae. These data do not help to distinguish between differing hypotheses on the
length and position of the raphe in the earliest raphe-bearing diatom.

The family Eunotiaceae was originally proposed by Kiitzing (1844:32) for those diatoms asym-
metrical about the longitudinal axis, with convex dorsal and concave ventral margins, and striae ex-
tending across the entire face of the valve. Today, we understand the family to contain those raphid
diatoms that also possess rimoportulae (Krammer and Lange-Bertalot 1991). The family is almost ex-
clusively freshwater. Included in the group are Eunotia Ehrenberg (which, in a broad sense, also in-
cludes species formerly separated into the genera Semiorbis Patrick and Desmogonium), Actinella
Lewis, Peronia Brébisson and Arnott, and Amphicampa Ehrenberg. Peronia has been reassigned to a
new family in which it is the only member (Round et al. 1990), but it is clearly allied to the genera men-
tioned above. Species of the Eunotiaceae are most diverse in dystrophic or acid waters.

Members of the Eunotiaceae are thought to have a unique systematic position, because of their
suite of characters, including both primitive (i.e., rimoportulae) and derived (i.e., raphe system) fea-
tures. Because both primitive and derived features are found in all members of the group (except some
highly derived forms that have secondarily lost a raphe system, see Kociolek and Rhode 1998), mem-
bers of the Eunotiaceae have been thought to represent early archetypes in the development of the
raphe system (Berg 1948). Some (e.g., Kolbe 1956; Geissler and Gerloff 1963) have speculated the
raphe evolved through enlargement of the rimoportulae along the ventral margin, by way of small
raphe-like slits (with taxa like Amphicampa proposed as intermediates between araphid and raphid di-
atoms). Hustedt (1952), in examining a wide range of eunotioid diatoms from the Amazon basin, be-
lieved the raphe evolved in a form similar to the genus Peronia (i.e., raphe elongated on the valve face
but without a central nodule). In Hustedt’s view, most of the Eunotiaceae represent a lineage off the
main evolutionary track towards the Naviculaceae. Simonsen (1979) aligned the Eunotiaceae with
one group of araphid diatoms, off the line in which the naviculacean raphe evolved. One implication
of Simonsen’s dendrogram is that the raphe in the Eunotiaceae is not homologous with the raphe sys-
tem of the naviculoids.

1]

12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2

In this report, I present light and scanning electron microscope observations on some
Eunotiaceae species that have been suggested by Hustedt (1952) and Kolbe (1956) to represent impor-
tant links in the evolution of the raphe system. Ultrastructural features are described and compared to
other members of the family and to other raphid diatom groups.

MATERIALS AND METHODS

Light and scanning electron microscope observations were conducted on several taxa within the
Eunotiaceae thought critical to understanding the early evolution of the raphe system. The taxa exam-
ined and material on which the observations were made include Eunotia curvula Hustedt (samples
AM 1027, AM 1028 from the Hustedt Collection), E. synedraeformis Hustedt (samples AM 1027,
AM 1028 from the Hustedt Collection), E. conversa Hustedt (samples AM 1027, AM 1030, AM 2216
from the Hustedt Collection),

Amphicampa eruca Ehrenberg (sample 607663 of the Diatom Collection, CAS) and Peronia
brasiliensis Hustedt (samples AM 1027, AM 1028, AM 2216 from the Hustedt Collection).

Hustedt material includes:

Sample AM 1027: “Brasilien. Lago Jurucui, an Pflanzen. 25.10.47. Braun, 252.”

Sample AM 1028: “Brasilien. Lago Jurucui, Siidufer, an Ast. 5.11.47. Braun, 253.”

Sample AM 1030: “Brasilien. Lago Jurucui, tiberschwemmte Campos, an Gr sern.
25.6.48. Braun 2548.”

Sample AM 2216: “Igarape do Tento. Uferschlick. 22.11.47. Braun 209.b.”

CAS sample 607663: “USA, California, Mendocino Co., Lake Mendocino, drying inlet to
lake. 28 Jun. 1995. coll. J. P. Kociolek.”

Material was cleaned in nitric acid, rinsed in distilled water, settled until neutral, and air-dried
onto coverslips. Coverslips containing the dried material were attached to aluminum stubs and sput-
ter-coated with approximately 20 nm of gold-palladium. The coated material was viewed on a Hitachi
S-520 SEM at an operating voltage of 10 kV.

RESULTS

Eunotia curvula Hustedt
Figures 1-4; Hustedt 1952, fig. 24; Simonsen 1987, pl. 568, figs. 1—5

DESCRIPTION. — Valves linear with margins nearly parallel, apices rounded, with convex dorsal
margin and concave ventral margin, 150-500 um long, 4.5—7.0 um broad. Raphe short, j-shaped,
curves from near the mantle onto the valve face, axial area distinct, striae 21/ 10 um, punctate.
Rimoportula present at each valve pole.

SEM OBSERVATIONS. — Round areolae are occluded externally. The raphe is hook- or j-shaped,
and has a large dilated distal end and undistinguished proximal end. A single rimoportula opening is
found at each valve terminus. The axial area extends the length of the valve. The valve is covered with
small siliceous nodules. Internally the raphe slit is bordered on both sides by a small ridge of silica.
The raphe runs from the proximal end, positioned on the valve face, towards the mantle, and recurves
towards the face, terminating in a prominent helictoglossa. A rimoportula is positioned at the valve
face-mantle junction.

KOCIOLEK: VALVE ULTRASTRUCTURE OF SOME EUNOTIACEAE 13

FicurES 1-4. Eunotia curvula, SEM. (1—2) External views at the apices. J-shaped raphe has dilated distal ends terminating
on valve face. Rimoportulae are positioned at the valve face-mantle junction. Valve face is covered with scattered siliceous
nodules. (3—4) Internal views at the apices. Raphe branches are restricted to ventral margin, with a prominent helictoglossa at
each pole. The raphe is bordered by a thin, elevated ridge. Rimoportulae are located near helictoglossa. Striae are interrupted
near the center of the valve.

Eunotia synedraeformis Hustedt
Figures 5-10; Hustedt 1952, fig. 8; Simonsen 1987, pl. 566, figs. 1-8

DESCRIPTION. — Valves straight or nearly so, with parallel margins, apices appear slightly swol-
len and broadly rounded, obtuse, length 235-525 um, breadth 9-10 tm. Raphe j- or hook-shaped, re-
curved on the valve face, axial area runs the length of the valve, striae punctate, with puncta near the
axial area more coarse and irregularly arranged within the striae. Striae number 18—22/ 10 um.

SEM OBSERVATIONS. — Rounded to nearly rectangular areolae are occluded externally. The
hook-shaped raphe is dilated at both the proximal end (positioned close to the valve mantle) and distal
end (positioned in the middle of the valve face). Striae are interrupted along the apical axis by a
hyaline area running the length of the valve. A narrowly elliptical, prominent opening of the
rimoportula is located at each pole and is positioned at the valve face-mantle junction. Internally,
areolae become more distantly spaced as they approach the center of the valve. Occlusions are lacking
internally. The raphe slit has a slight ridge bordering it as it runs from the proximal end on the valve

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2

FIGURES 5—10. Eunotia synedraeformis, SEM. (5—6) External views at the apices. Prominent J-shaped raphe with dilated dis-
tal ends recurved towards center of valve. Rimoportula opening is visible. Occluded areolae appear sunken. (7-8) Internal
views at the apices. Raphe opening is restricted to ventral margin, bordered by a thin, elevated ridge, and ends in large
helictoglossae. Rimoportulae are found at both poles. (9) Center of the valve. External view showing interruption of striae
forming central sternum. (10) Center of the valve. Internal view showing disruption of striae towards center.

KOCIOLEK: VALVE ULTRASTRUCTURE OF SOME EUNOTIACEAE 15

face towards the mantle and recurves towards the valve center at the distal end. The raphe terminates
in a prominent helictoglossa at its distal end. Located near the helictogossae at the valve-mantle junc-
tion are well-developed rimoportulae.

Eunotia conversa Hustedt
Figures 11—16; Hustedt 1952, fig. 25; Simonsen 1987, pl. 565, figs. 1-8

DESCRIPTION. — Valves narrow, straight for most of the length of the valve, bent about the apical
axis, ends narrowly rounded. 50—120 um long, 2.0—3.5 um broad. Raphe j-shaped, but difficult to dis-
cern. Axial area present but indistinct. Striae about 28—30/10 um.

SEM OBSERVATIONS. — Internally the raphe slit is located in a raised central rib and extends
along the center of the valve. The raphe terminates in a helictoglossa. A rimoportula is found on the
face at the valve terminus at only one end of each valve. An internally-raised ridge is found positioned
towards the ventral margin of the valve. The ridge extends about 1/3 of the length of the raphe. Striae
are interrupted by a wide unornamented axial area near the center of the valve . Externally, the rela-
tively long raphe slit curves and continues back towards the center of the valve. The shorter portion of
the raphe branch does not perforate the valve. A prominent opening of the single rmoportula is found
at the junction of the valve face and mantle. Areolae appear as small, round openings internally.

Peronia brasiliensis Hustedt
Figures 17—20; Hustedt 1952, figs. 1, 2; Simonsen 1987, pl. 562, figs. 1—7

DESCRIPTION. — Valves narrow, asymmetrical about the transapical axis and have rounded
poles. Length 15—30 um, breadth 3—S um. The raphe branches are short, restricted to the apices, and
positioned in the middle of the valve face. Striae are parallel, slightly radiate at the poles,
20—30/10 um.

SEM OBSERVATIONS. — Internally, the centrally-positioned, short raphe slits extend from the
proximal ends and terminate as prominent helictoglossae. A small, relatively inconspicuous
rimoportula is located at one pole only, at the valve terminus. A central nodule is lacking. Between the
proximal raphe ends is located an unornamented axial area. The round puncta appear to lack occlu-
sions.

Amphicampa eruca Ehrenberg
Figures 21—35

DESCRIPTION. — Valves19—64 ym long, 9-11 um broad. Both the dorsal and ventral margins
are undulate; the dorsal margin has 3—7 undulations, whereas the ventral margin has |—6 undulations.
Striae are distinctly punctate, and the discontinuity across the striae is closer to the ventral margin.
Small raphe slits are indistinct but visible near the valve terminus close to the ventral margin.

SEM OBSERVATIONS. — The external valve face is covered by striae composed of rounded
areolae. Striae are interrupted towards the ventral margin on the valve face. Distinct breaks in the
striae also occur around the margin of the valve. Long and short striae alternate on the valve mantle.
Short raphe branches of variable shape are positoned close to the valve face-mantlejunction. Spines
are lacking. Internally, raphe branch length and helictoglossa size are quite variable. A rimoportula is
positoned at each end of the valve. The position of the rimoportulae is variable; they may be placed at
the valve terminus, at the terminus/dorsal margin junction, or along the dorsal margin.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2

FiGuRES 1 1—16. Eunotia conversa, SEM. (11) External view. Elongate raphe sharply recurved towards valve center. Single
rimoportula opening is visible. (12) External view. Valve apex with prominent raphe branch and occluded areolae. Fics. 13-16.
Internal views. (13—14) Valve apices of same valve showing elongate raphe slit bordered by thin, elevated ridge. One end (Fig.
13) has arimoportula. Linear swelling indicates where external groove of raphe is located. (15) Proximal raphe end undifferen-
tiated, with lack of a central nodule. (16) Center of valve showing striae interrupted in middle of valve.

KOCIOLEK: VALVE ULTRASTRUCTURE OF SOME EUNOTIACEAE

FIGURES 17-20. Peronia brasiliensis, SEM, internal views. (17-18) Headpole (Fig. 17) and footpole (Fig. 18) of the same
valve. Small raphe branches are restricted to apices. Helictoglossae are prominent. Single rimoportule is located at headpole.
(19-20) Headpole (Fig. 19) and footpole (Fig. 20) of the same valve. Headpole shows striae bordering raphe to be more dense.
Striae are composed of round puncta. Rimoportule is located near helictoglossa at headpole. Central nodule is lacking.

18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2

FIGURES 21-29. Amphicampa eruca, LM. Valve views showing variation in outline and size diminution. Scale bar is 10 pm
for all figures.

DISCUSSION

Observations presented here provide additional evidence of considerable variation in raphe
structure in the Eunotiaceae. This variation includes: (1) a short raphe system without a central ster-
num, restricted to the mantle (in Amphicampa); (2) a longer raphe system without a central sternum ei-
ther restricted to or predominantly on the mantle (in Eunotia and Actinella; Round et al. 1990;
Lange-Bertalot 1995; Kociolek et al. 1997); (3) an even longer, highly curved raphe mostly on the
valve face with a central sternum (E. conversa, E. curvula, E. synedraeformis); and (4) a straight raphe
centrally positioned on the valve face with a central sternum (Peronia brasiliensis).

The observations, which are based on raphe features, suggest there is variation within a genus, as
well as within the family Eunotiaceae. For example, Round, Crawford, and Mann (1990:459, fig. m)
illustrated what looks like a simple central nodule ina species of Peronia, and the lack ofa central nod-
ule in P. brasiliensis might suggest that Peronia is a non-monophyletic group. Clearly, with presence
of rimoportulae being a primitive feature and presence of a raphe diagnosing the entire group of raphid
diatoms, this suite of features cannot be used to diagnose the Eunotiaceae as has been done in the past
(e.g., Krammer and Lange-Bertalot 1991). It is therefore possible that other rimoportula-bearing
raphid diatoms do not necessarily belong to the Eunotiaceae (in either the strict or broad sense); an ex-
ample of this is the genus Eunophora, which may be more closely aligned to Amphora and its relatives

KOCIOLEK: VALVE ULTRASTRUCTURE OF SOME EUNOTIACEAE 19

FiGures 30-35. Amphicampa eruca, SEM. (30-31) External views at the apices. Both figures show small raphe branches at
ventral margin. Discontinuity in striae is located towards ventral margin. Striae are composed of round, unoccluded areolae.
Spines are lacking. (32-33) Internal views at apices. Opposite ends of same valve showing presence of rimoportulae located on

valve mantle (arrows). Raphe branches are small, with indistinct helictoglossae. (3435) Internal views at apices. Apices of dif-
ferent cells showing variation in position of rimportulae (arrows). Raphe branches and helictoglossae are noticeable.

20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2

(Vyverman et al. 1998). Presence of a central nodule in some Peronia species would lend support to
the creation of the Peroniaceae (Round et al. 1990), since a central nodule is absent in the members of
the Eunotiaceae.

Eunotia sensu stricto appears to be composed of many disparate morphological subgroups,
whose relationships within the Eunotiaceae require further research. Hustedt (1926), for example,
suggested the earliest branch within Eunotia may be composed of the bilunaris group. Krammer and
Lange-Bertalot (1991) have documented with SEM the ultrastructure of E. bilunaris and its allies,
which have raphe and valve construction similar to E. curvula and E. synedraeformis. This similarity
may suggest that the bilunaris group may have evolved separately from other Eunotia groups. L
ange-Bertalot’s (1995) E. weisingii may be a transitional form between E. conversa and the bilunaris
group. Variation among other Eunotia taxa has been documented by Krammer and Lange-Bertalot
(1991) and Lange-Bertalot (1995). An implication of these observations is that Eunotia may not nec-
essarily be amonophyletic group. More work is necessary to resolve the finer relationships within the
Eunotiaceae.

While we may now understand the variation in raphe expression to be greater than previously ap-
preciated, our observations do not yet speak to the question of the evolution of the raphe system from
araphid ancestors. Kolbe’s (1956) hypothesis of the earliest raphid-bearing diatom being similar to
Amphicampa (supported by the work of Mann 1984), then evolving through forms with progressively
longer raphe systems until the naviculoid type is reached, is consistent with our observations. How-
ever, equally plausible is the idea that the earliest raphe system was fully developed (similar to that in
Peronia brasiliensis, Hustedt 1952; see also Hustedt 1935), and that the rest of the eunotioid forms
represent degeneration, with Amphicampa showing the greatest expression of raphe reduction.
Kociolek and Rhode (1998) have suggested raphe reduction occurred elsewhere within the
rhaphidiod line. In the Actinel/la lineage from Madagascar, a series of raphe types were observed, and
suggested to represent reduction of a typical Actinella raphe system to a simple slit without a
helictoglossa (Kociolek and Rhode 1998). Taxa such as Falcula Voigt, shown to have a slit positioned
close to the ventral margin may reflect a degenerated (but derived) condition of the raphe, rather than a
primitive raphe condition, as argued by Geissler and Gerloff (1963). Given the diversity within
Amphicampa (Ehrenberg 1854) and the closely allied genera Ophidocampa and Heterocampa
(Ehrenberg 1870), both of which have not been investigated since the taxa were described originally,
the relationship of Amphicampa and its close allies within the Eunotiaceae needs further investiga-
tion. Also, examination of original Ehrenberg material to confirm the identity of Amphicampa eruca
is crucial to confirming the present results.

Further observations on other naviculoid forms, including Rouxia, Amphipleura and Frustulia
among others, as well as other raphe- and rimoportula-bearing taxa (such as Eunophora) will be criti-
cal in developing a formal phylogenetic hypothesis on the origin and early evolution of the raphe sys-
tem.

ACKNOWLEDGMENTS

Sarah Spaulding, CAS, graciously provided technical assistance. Discussions with participants
of the XIV North American Diatom Symposium (1997) about these findings were particularly help-
ful. Reviews of this manuscript by S. Spaulding, E. Fourtanier, R. Jahn, D. Williams and the research
group in the laboratory of E. F. Stoermer were also helpful.

KOCIOLEK: VALVE ULTRASTRUCTURE OF SOME EUNOTIACEAE 21

LITERATURE CITED

BERG, A. 1948. Observations on the development of the Eunotia-raphe. Arch. Botanik. 33A:1—10.

EHRENBERG, C. G. 1854. Mikrogeologie. Das Erden und Felsen schaffende Wirken des unsichtbar Kleinen
selbstanigen Lebens auf der Erde. Leopold Voss, Leipzig. 374 pp.

. 1870. Uber machtige Gebirgs-Schichten vorherrschend aus mikroskopischen Bacillarien unter und bei
der Stadt Mexiko. Physik. Abhandl. kénig. Akad. Wiss. Berlin. 1869:1—66.

GEISSLER, U. AND J. GERLOFF. 1963. Elektronenmikroskopische Beitrage zur Phylogenie der Diatomeenrhaphe.
Nova Hedwigia 6:339-352.

HASLE, G. R. 1973. The “mucilage pore” of pennate diatoms. Beih. Nova Hedwigia 45:167—186.

HUSTEDT, F. 1926. Untersuchungen tiber den Bau der Diatomeen. I. Raphe und Gallertporen der Eunotioideae.
Ber. Deutsche Bot. Gesell. 44:142—150.

. 1935. Untersuchungen tiber den Bau der Diatomeen. XII. Ber. Deutsche Bot. Gesell. 53:246-264.

. 1952. Neue und wenig bekannte Diatomeen III. Phylogenetische Variationen bei den rhaphidioiden
Diatomeen. Ber. Deutsche Bot. Gesell. 65:133-144.

KOCIOLEK J. P. AND K. RHODE. 1998. Raphe vestiges in ““Asterionella” species from Madagascar: evidence for a
polyphyletic origin of the araphid diatoms? Crypto. Algol. 19:57—74.

KOCIOLEK J. P., K. RHODE, AND D. M. WILLIAMS. 1997. Taxonomy, ultrastructure and biogeography of the
Actinella punctata species complex (Bacillariophyta: Eunotiaceae). Nova Hedwigia 65:177—193.

KOLBE, R. W. 1956. Zur Phylogenie des Raphe-Organs der Diatomeen: Eunotia (Amphicampa) eruca Ehr. Bot.
Notiser. 109:91—97.

KRAMMER, K. AND H. LANGE-BERTALOT. 1991. Bacillariophyceae 2/3. Centrales, Fragilariaceae, Eunotiaceae.
In Siisswasserflora von Mitteleuropa, H. Ettl, J. Gerloff, H. Heynig and D. Mollenhauer, eds. Gustav Fisher,
Stuttgart. 576 pp.

KUTZING, F. T. 1844. Die Kieselschaligen Bacillareen oder Diatomeen. Nordhausen. 152 pp.

LANGE-BERTALOT, H. 1995. 85 Neue Taxa und iber 100 weitere neu definierte Taxa erganzend zur
Susswasserflora von Mitteleuropa Vol. 2/14. Bibl. Diatomol. 27:1-454.

MANN, D. G. 1984. An ontogenetic approach to diatom systematics. Pp. 1 13—144 in Proceedings of the 7th Inter-
national Diatom Symposium, D.G. Mann, ed.; 1982 August 22-27; Philadelphia, PA. O. Koeltz,
Koenigstein. 541 pp.

ROUND F. E., R. M. CRAWFORD, AND D. G. MANN. 1990. The diatoms. Morphology and biology of the genera.
Cambridge University Press, Cambridge, U. K. 747 pp.

SIMONSEN, R. 1979. The diatom system: ideas on phylogeny. Bacillaria 2:9—71.

. 1987. Atlas and catalogue of the diatom types of Friedrich Hustedt. 3 Vols. J. Cramer, Berlin.

VYVERMAN, W., K. SABBE, D. G. MANN, C. KILROY, R. VYVERMAN, K. VANHOUTTE, AND D. HODGSON. 1998.
Eunophora gen. nov. (Bacillariophyta) from Tasmania and New Zealand: description and comparison with
Eunotia and amphoroid diatoms. European J. of Phycology 33:95—111.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 3, pp. 23-31, 3 figs., 2 tables. February 23, 2000

Silurian Polyplacophora and Rostroconchia (Mollusca) from
Northern California

by
Richard D. Hoare MAP :
Department of Geology, Bowling Green State University = © <
Bowling Green, Ohio 43403

Silicified polyplacophorans from the Gazelle Formation in California represent the second
known Silurian (Ludlovian) occurrence of this class in North America. New taxa are
Paleochiton siskiyouensis sp. noy. and Thairoplax merriami sp. nov. Eusphairica distubula
gen. et sp. nov. is the first Silurian rostroconch identified in the United States and extends
the range of the family Eopteriidae Miller, 1889, from the Middle Ordovician into the Up-
per Silurian.

The late C. W. Merriam (1972), U. S. Geological Survey, collected and processed samples of
limestone cobbles with silicified fossils from a conglomeratic zone of the Silurian Gazelle Formation
exposed in Siskiyou County, California, in 1951, and noted the presence of amphineuran plates. Some
of the specimens obtained were placed in the collections of the California Academy of Sciences. Peter
U. Rodda brought this polyplacophoran fauna to my attention.

Preservation of the silicified plates range from poor to excellent. Many are imcomplete and most
have additional siliceous deposits adhering to them. All are intermediate plates with the exception of
two tail plates. Head plates were not found among the more than 50 specimens.

A single specimen of a rostroconch is present with a portion of a chiton plate attached to it. The
importance of this rostroconch occurrence necessitated the removal of as much of the plate and other
adhering material as possible to allow better photographic representation of the specimen.

Based on the rugose coral fauna described by Merriam (1972), and the accompanying fauna of
brachiopods, gastropods, and trilobites, the Gazelle Formation is mainly Late Silurian (Ludlovian) in
age. Portions of the formation in this region may be slightly older or younger than this (Merriam
1972:23):

The collection locality in the Gazelle Formation, which represents the type locality for all of the
new taxa herein, is 2 km southeast of Parker Ranch, East Fork of the Scott River, SW 4 sec. 29, T41N,
R7W, Siskiyou County, California, Etna quadrangle, USGS locality M1027.

PREVIOUS STUDIES

European Silurian polyplacophorans have been studied by Salter (in M’Coy 1846), de Koninck
(1857; translated by Bailey 1860), Barrande (1867), Davidson and King (1874), Lindstrom (1884),
Woodward (1885), Etheridge (1897), Couper Reed (1911), Bergenhayn (1943, 1955), and Cherns
(1998a, 1998b, 1999).

Other than the noted presence by Merriam (1972:15), the only other account of North American
Silurian polyplacophorans was by Kluessendorf (1987), from Wisconsin, Illinois, and Iowa. Col-
lected from carbonate buildups, these specimens are preserved as impressions of the ventral surface.

23

24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 3

Kluessendorf erected, Chelodes raaschi sp. nov. and Hawthorneachiton lowenstami gen. et. sp. nov.
Other specimens were described as morphotypes. One of the latter, Morphotype H (Kluessendorf
1987, fig. 10), represents a turrileped plate.

SPECIES DESCRIPTIONS

Class Polyplacophora de Blainville, 1816
Subclass Paleoloricata Bergenhayn, 1955
Order Chelodida Bergenhayn, 1955
Suborder Chelodina Bergenhayn, 1955
Family Gotlandochitonidae Bergenhayn, 1955
Genus Paleochiton Smith in Smith and Toomey, 1964

Paleochiton siskiyouensis sp. nov.
Fig. |

TYPE MATERIAL. — HOLOTYPE: CAS 68395, PARATYPES: CAS 68396.01—68396.18.

DIAGNOSIS. — Intermediate plates strongly arched; shallow to deep jugal sinus; apical area
short, bordering nearly straight to slightly produced posterior margin.

DESCRIPTION. — Intermediate plates strongly arched, subrectangular in dorsal view; jugal area
rounded; side slopes flatly convex; posterior margin straight to slightly mucronate; lateral margins
straight to slightly convex, nearly parallel; anterior margin with deep jugal sinus; jugal angle 83°-89°;
apical area short, 1.4 mm in length, straight to flatly V-shaped; shell material thick medially, thinning
laterally; surface smooth.

Tail plate wedge-shaped in dorsal view; posterior margin pointed, curved dorsally; anterior mar-
gin flatly convex; jugal area broadly rounded, side slopes flat.

MEASUREMENTS. — See Table 1.

DISCUSSIONS. — Paleochiton siskiyouensis differs from the Ordovician P. kindbladensis Smith
in Smith and Toomey, 1964, by having a larger length:width ratio 1.26:1.14, a smaller jugal angle
86°—114°, and smaller valve size. Kluessendorf’s Morphotype A (1987:439, fig. 4, Pl. 1, fig. 3) may
be a Paleochiton but has a larger jugal angle, 100°—120°, and is a larger plate. The anterior margin is
unknown.

ETYMOLOGY. — Named for Siskiyou County, California.

Genus Thairoplax Cherns, 1998b

Thairoplax merriami sp. nov.
Figs. 2A—N

TYPE MATERIAL. — HOLOTYPE: CAS 68397, PARATYPES: CAS 68398.01—68398.10.

DIAGNOSIS. — Intermediate plates strongly mucronate, strongly arched; anterior margin with
well-developed jugal sinus; apical area large, V-shaped.

DESCRIPTION. — Intermediate plates relatively thin, strongly mucronate with slightly convex
posterolateral margins converging on apex; highly arched with narrow jugal area; jugal angle
95°—106°; side slopes flat, trapezoidal in outline; anterolateral margins often converging slightly an-
teriorly, curving narrowly into anterior margin; jugal sinus prominent, fairly deep; apical area large,
2.3 mm long, V-shaped, approximately one-third of total length at midline; shell material moderately
thick medially, thinning laterally; surface smooth.

HOARE: NEW SILURIAN MOLLUSKS FROM CALIFORNIA 25

Q

FIGURE 1. Paleochiton siskiyouensis sp. nov. A, B. Intermediate plate, ventral and posterior views (CAS 68396.01); C—E. In-
termediate plate, dorsal, ventral, and left lateral views (CAS 68396.02): FI. Intermediate plate, dorsal, ventral, left lateral, and
anterior views (CAS 68396.03); J, K. Holotype, intermediate plate, dorsal and ventral views (CAS 68395): L, M. Intermediate
plate, dorsal and anterior views (CAS 68396.04); N. Intermediate plate, dorsal view (CAS 68396.06): O. P. Intermediate plate
with a second plate beneath it, right lateral and dorsal views (CAS 68396.08); QT. Tail plate, dorsal, ventral, left lateral, and
anterior views (CAS 68396.05). All figures x 5.

26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 3

TABLE |. Measurements (in mm) of Paleochiton siskivouensis sp. NOV.

CAS Collection No. Length Width Height Apical area
68395* 13 5.6 3.0 12
68396.01 ees a7, 3 12
68396.02 6.0 5.8 1.6 2
68396.03 ~ 4.8 3.0 -
68396.04 - 6.4 3.8 1.0
68396.05 6.9 4.1 3.4 -
68396.06 ~ SO LES -
68396.07 5.4 5.0 3.07 [2
68396.08 SB 5.0 355 1.6
68396.09 5.9 4.9 2.6 ~
68396.10 a3 5.4 3.4 3
68396.11 7.0 5.8 3.0 12
68396.12 5.8t 5.0 DS 1.1
68396.13 6.57 6.0 Bul -
68396.14 6.0 SE) ul -
68396.15 48 5.6 2.6 1.4
68396.16 55 5.0 2.8 0.7
68396.17 6.3 3.4 2.4 0.8
68396.18 5.8 5.0 23 -
* Holotype

+ Estimated

Tail plate with well-developed jugal sinus; anterior lateral margins subparallel; posterolateral
margins converging to posterior apex; not as strongly arched as intermediate plates; surface smooth.
Head plate unknown.

MEASUREMENTS. — See Table 2.

DISCUSSION. — The genus 7hairoplax was established by Cherns (1998b:946) for Silurian spe-
cies from Gotland, Sweden, one of which had been previously placed in Gotlandochiton Bergenhayn,
1955. Thairoplax differs from the latter genus by its V-shaped apical area, more acute apex, and
greater length:width ratio of intermediate plates. The primary difference between 7. merriami and T.
pelta Cherns, 1998b, the type species, is the larger, V-shaped apical area and converging anterordorsal
margin in 7. merriami. Thairoplax differs from Chelodes Davidson and King, 1874, by having thinner
shell material, a sharp jugal flexure, and flat side slopes.

ETYMOLOGY. — Named for the late Charles W. Merriam, U. S. Geological Survey, who col-
lected the specimens.

Family Mattheviidae Walcott, 1886

Genus and species A
Figs. 20-T

MATERIAL. — CAS 68399.01, CAS 68399.02.

DESCRIPTION. — Intermediate plates subquadrate, thick, broadly arched; little to no distinction
between jugum and side slopes; anterior margin with well-developed jugal sinus; posterior margin
straight to slightly mucronate; posteror lateral margins converging posteriorly; apical area very short;
surface smooth. Head and tail plates unknown.

HOARE: NEW SILURIAN MOLLUSKS FROM CALIFORNIA Ped

FIGURE 2. A-N. Thairoplax merriami sp. nov. A. Intermediate plate, dorsal view (CAS 68398.01); B. Intermediate plate,
ventral view (CAS 68398.02); C, D. Intermediate plate, posterior and dorsal views (CAS 68398.03); E. Intermediate plate, ven-
tral view (CAS 68398.04): F. Intermediate plate, ventral view (CAS 68398.05); G, H. Intermediate plate, ventral and right lat-
eral views (CAS 68398.06); I-K. Holotype, intermediate plate, dorsal, ventral, and anterior views (CAS 68397): L-N.
Intermediate plate, dorsal, ventral, and left lateral views (CAS 68398.07). O-T. Genus et sp. A. O-Q. Intermediate plate, dorsal,
anterior, and ventral views (CAS 68399.01); R—T. Intermediate plate, dorsal, left lateral, and ventral views (CAS 68399.02).
All figures x 5.

28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 3

TABLE 2. Measurements (in mm) of Thairoplax merriami sp. nov.

CAS Collection No. Length Width Height Apical area
68397* 8.5 5.8 2.8 pA
68398.01 7.6 5G 3u1 3.0
68398.02 8.8 52 2.4 2.9
68398.03 Sy) 44 22 2.0
68398.04 th 5.07 23 2.4
68398.05 6.6 6.07 3.07 2.6
68398 .06 6.4 5.07 32 1.3
68398 .07 Ve 4.5 25 20
68398.08 7.4 6.2 3.0 ~
68398.09 ep) 5.0 am 20
68398.10 6.9 4.8 3.4 -

* Holotype
¥* Estimated

MEASUREMENTS. — Two plates range in length, 5.6—7.2 mm: in width, 6.9—6.8 mm: in height.
3.5-3.1 mm: in length of apical area. 0.5—0.6 mm.

DISCUSSION. — Two intermediate plates in the collection differ from both Paleochiton
siskivouensis and Thairoplax merriamiin having a transversely thickened ridge on the ventral surface
somewhat similar to that on A/astega lira Cherns, 1998b, from the Silurian of Sweden, but the latter
taxon is more strongly arched and mucronate than the California specimens. The genus Chelodes
Davidson and King, 1874, is similar in plate thickness and jugal sinus but is more mucronate, with a
large apical area. Kindbladochiton Van Belle, 1975, and /voechiton Bergenhayn, 1955, do not have as
great a shell thickness and are more strongly arched than the specimens described here, but they may
represent a species of the latter genus.

Class Rostroconchia Pojeta. Runnegar, Morris, and Newell, 1972
Order Conocardioida Neumayr. 1891
Superfamily Eopteriacea Miller, 1889
Family Eopteriidae Miller, 1889

Genus Eusphairica gen. nov.

TYPE SPECIES. — Eusphairica distubula sp. nov.

DIAGNOSIS. — Body subcircular in cross section: snout short: rostrum lacking: gape extending
from just ventral of rostral opening to anterior end: posterior clefts present.

DESCRIPTION. — See Eusphairica distubula sp. nov.

DISCUSSION. — Eusphairica is monotypic. It differs from the Ordovician genera Eopteria Bill-
ings, 1865, and Euchasuna Billings, 1865, by its more tumid body shape, lack of a rudimentary ros-
trum, and gape not extending to the rostral opening. Both Wanwanella and Wanwanoidea, both by
Kobayashi, 1933, as illustrated by Pojeta and Runnegar (1970), are narrower forms with less distant
snouts than Eusphairica.

ETYMOLOGY. — Greek ew meaning primitive, good; plus sphairica, globular.

HOARE: NEW SILURIAN MOLLUSKS FROM CALIFORNIA 29

Eusphairica distubula sp. nov.
Fig. 3

TYPE MATERIAL. — HOLOTYPE: CAS68400.

DIAGNOSIS. — Same as genus.

DESCRIPTION. — Body small, subcircular in cross section: dorsal portion of posterior face ex-
tending posteriorly, slanted ventrally, curving sharply just below rostral opening and slanting anteri-
orly; rostrum not present: dorsal margin straight: beak slightly posterior to midlength: ventral margin
of body straight, curving concavely into short snout: snout with keyhole gape anteriorly: ventral gape
narrow, extending posteriorly to just ventral of rostral opening: body with marginal denticles in gape:
area around rostral opening smooth or with nonpreserved nbs: central portion of face with five to six
fine ribs: four coarser radial nbs at juncture of posterior face and body: 13 or more finer radial ribs on
body: anterior portion of snout may lack radial ribs: comarginal growth lines present ventrally on
body and snout: internal features not observed.

MEASUREMENTS. — Length 7.0 mm: width 5.2 mm: height 5.2 mm: width of anterior end
1.7 mm: length of ventral gape 6.2 mm: diameter of rostral opening 0.8 mm.

DISCUSSION. — The subspheroidal shape of the body, the slightly angular projection of the ex-
tended posterior face, and the gape ending ventral to the rostral opening are diagnostic of Eusphairica
distubula. Eopteria struszi Pojeta, Gilbert-Tomlinson, and Shergold, 1977, described from the Lower
Ordovician of Australia, is somewhat similar to Eusphairica distubula but differs in being narrower,
having coarser radial ribs, a rudimentary rostrum, and a gape extending into the rostrum. Johnson and
Chatterton (1983) described nine new species representing five genera of rostroconchs from the Mid-
dle Silurian of the Northwest Territories, Canada. None of these beautifully preserved, silicified spec-

FiGURE 3. Eusphairica distubula gen. et sp. nov. (CAS 68400). A. Right lateral view; B. Dorsal view; C. Ventral view; D.
Anterior view; E. Ventroposterior view; F. Posterior view. All figures x 7.

30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 3

imens have the characters of the posterior face, ventral gape, and arrangement of ribs as are present on
E. distubula.
ETYMOLOGY. — Latin dis meaning without; plus twbulus, tube (rostrum).

ACKNOWLEDGMENTS

I wish to thank P. U. Rodda. former Chairman of the Department of Geology. California Acad-
emy of Sciences, who notified me of the presence of the collectionand J. DeMouthe, California Acad-
emy of Sciences, for making the specimens available for study. J. Thomas Dutro, Jr.. U. S. Geological
Survey. provided the source of information concerning the collection locality.

LITERATURE CITED

BAILEY, W. H. 1860. Observations on two new species of Chiton from the Upper Silurian “Wenlock Limestone”
of Dudley. Annales and Magazine of Natural History 6(32):91—98. (An English translation of de Koninck,
1857).

BARRANDE, J. 1867. Systeme Silurien de centre de la Boheme. Premiere partie: recherches paléontologiques,
tom. 3. Classe des Mollusques. Order des Ptéropodes. Prague and Paris. 179 pp.

BERGENHAYN, J. R.M. 1943. Preliminary notes on the fossil polyplacophoras trom Sweden. Geologiska
Féreningens, I, Stockholms, Férhandlingar 65(3):297-303.

. 1955. Die Fossilien Schwedischen Loricaten nebst einer vorlaufigen Revision des Systems der ganzen
Klasse Loricata. Lund Universitet, Arsskrift. N.F. Avd. 2, 51(8); Kungliga Fysiografiska Sallskapets,
Handlingar, N.F, Avd. 2, 66(8):1-41.

BILtinas, E. 1865. Palaeozoic fossils, containing descriptions and figures of new or little known species of or-
ganic remains from the Silurian rocks, 1861—1865. Geological Survey of Canada, Volume 1, 426 pp.
BLAINVILLE, H. M. D. De. 1816. Prodrome d-une nouvelle distribution systématique du regne animal. Bulletin

Sciences Société Philomathigue de Paris 105—124.

CHERNS, L. 1998a. Chelodes and closely related Polyplacophora (Mollusca) from the Silurian of Gotland, Swe-
den. Palaeontology 41(3):545—5S73.

.1998b. Silurian polyplacophoran molluscs from Gotland, Sweden. Palaeontology 41(5):939—974.

1999. Silurian chitons as indicators of rocky shores and lowstand on Gotland, Sweden. Palaios
14(2):172-179.

COUPER REED, F. R. 1911. A new fossil from Girvan. The Geological Magazine 8(8):337-339.

DAVIDSON, T. AND W. KiNG. 1874. On the Trimerellidae,a Paleozoic family of the palliobranchsor Brachiopoda.
Quarterly Journal of the Geological Society of London 30(2): 124-172.

ETHERIDGE, JR., R. 1897. On the occurrence of the genus Chelodes Davidson and King, in the Upper Silurian of
New South Wales. Records of the Geological Survey of New South Wales 5(2):67—70.

JoHNsTON, D.I. AND B. D. E. CHATTERTON. 1983. Some silicified Middle Silurian rostroconchs (Mollusca)
from the Mackenzie Mountarins, N. W. T., Canada. Canadian Journal of Earth Science 20(5):844-858.

KLUESSENDORF, J. 1987. First report of Polyplacophora (Mollusca) from the Silurian of North America. Cana-
dian Journal of Earth Science 24:435-441.

KoBAYASHI, T. 1933. Faunal study of the Wanwanian (basal Ordovician) series with special notes on the
Ribeiridaeand the ellesmereiceroids. Tokyo Imperial University Faculty of Science Journal 3(7):249-328.

Koninck, M. L. DE. 1857. Sur deux nouvelles espéces siluriennes appartenant au genre Chiton. Bulletin de
1’ Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique 11:190—199.

LINDSTROM, G. 1884. On the Silunan Gastropoda and Pteropoda of Gotland. Kongliga Svenska
Vetenskaps-A kademiens Handlingar 19(6):48—52.

M’Coy, F. 1846. A synopsis of the Silurian fossils of Ireland. Privately issued in 1846 by R. Griffiths, Dublin.
207 pp.

MERRIAM, C. W. 1972. Silurian rugose corals of the Klamath Mountains Region, California. U. S. Geological
Survey Professional Paper 738, 50 pp.

MILLER, S. A. 1889. North American geology and paleontology for the use of amateurs, students and scientists.
Western Methodist Book Concern, Cincinnati. 664 pp.

HOARE: NEW SILURIAN MOLLUSKS FROM CALIFORNIA 31

NeuMAYR, M. 1891. Beitraége zu einer morphologischen Eintheilung der Bivalven. K. Akademie der
Wissenschaften zu Wien Denkschriften 58:701—801.

PosETA JR., J. AND B. RUNNEGAR. 1976. The paleontology of rostroconch mollusks and the early history of the
Phylum Mollusca. U. S. Geological Survey Professional Paper 968, 88 pp.

PoreTA JR., J., B. RUNNEGAR, N. J. MorRIS, AND N. D. NEWELL. 1972. Rostroconchia: a new class of bivalved
mollusks. Science 177 (4045 ):264—267.

PosETA JR., J., J. GILBERT-TOMLINSON, AND J. H. SHERGOLD. 1977. Cambrian and Ordovician rostroconch
molluscs from Northem Australia. Australian Bureau of Mineral Resources, Geology and Geophysics Bul-
letin 171, 54 pp.

SALTER, J. W. 1847. Description of a fossil Chiton from the Silurian rocks, with remarks on the fossil species of
the genus. Quarterly Journal of the Geological Society of London 3:48—52.

SMITH, A. G. AND D. F. Toomey. 1964. Chitons from the Kindblade Formation. Oklahoma Geological Survey
Circular 66, 41 pp.

VAN BELLE, R.A. 1975. Sur la classification der Polyplacophora: I. Introduction at classification du
Paleoloricata, avec la description de Kindbladochiton nom. nov. (pour Eochiton Smith, 1964). Informations
de la Societe Belge de Malacologie 5:121—145.

Wa corr, C. D. 1886. Studies on the Cambrian faunas of North America. U. S. Geological Survey Bulletin 30,
369 pp.

WooDWaRrbD, H. 1885. Ona new species of Helminthochiton from the Upper Bala (Silurian) of Girvan, Ayrshire.
Geological Magazine 2:352-355.

© CALIFORNIA ACADEMY OF SCIENCES. 2000
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 4, pp. 33-54, 9 figs. May 10, 2000

New Costa Rican and Panamanian Species of Miconia
(Melastomataceae: Miconieae)

by

Frank Almeda
Department of Botany, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Diagnoses, descriptions, and illustrations are presented for seven new Mesoamerican spe-
cies of Miconia (M. colliculosa and M. talamancensis from Costa Rica and Panama, M.
vestita from Costa Rica, and M. correae, M. crocata, M. jefensis, and M. morii from Pan-
ama). Distinguishing characters, distribution maps, citations of representative specimens,
and comparisons with probable relatives are provided for each species.

RESUMEN

Miconia, con mas de 1,000 especies descritas, es el género mas grande en la region de la
Flora Mesoamericana. Se describen siete especies nuevas de Miconia (M. colliculosa y M.
talamancensis de Costa Rica y Panama; M. vestita de Costa Rica; y M. correae, M. crocata,
M. jefensis y M. morii de Panama). Se proveen descripciones, ilustraciones, notas sobre
distribucién y fenologia para todas las especies. Ademas se presentan discusiones sobre las
afinidades entre las especies nuevas y mapas de distribucion.

In the course of preparing a floristic treatment of the Melastomataceae for Flora Mesoamericana
many new taxa have come to light. Several are from remote, little-collected regions while others are
locally abundant at sites that have received repeated visits from collectors during the past two de-
cades. Because work on this treatment has necessitated study of collections from well beyond the lim-
its of the flora area, it has been possible to definitively identify new taxa and gain a better
understanding of variation patterns in problematic species complexes over a broad geographic area.
Of the 37 genera and approximately 500 species of melastomes presently recorded for the
Mesoamerican region, 162 of the species are in the genus Miconia. Of this total, 126 species of
Miconia occur in Costa Rica and Panama. It comes as no surprise, therefore, that the lion’s share of
novelties continues to come from these two countries, both of which are important centers of
biodiversity in northern Latin America.

SPECIES DESCRIPTION

Miconia colliculosa Almeda, sp. nov.
Fig. 1

TYPE. — COSTA RICA. Limon: Canton de Talamanca Amubri. Camino entre Amubri y Soki.
Siguiendo el Rio Nabri hacia Alto Soki, 9°29'50”"N, 82°59'10"W, elev. 150 m, 3 Jul. 1989, Herrera
3129 (holotype: CAS!; isotypes: CR!, INB!, MEXU! MO).

33

34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

Section Amblyarrhena. Frutex vel arbuscula 2-6 m. Ramuli teretes sicut petioli foliorum subtus
venae primariae inflorescentia hypanthiaque dense pilis penicillato-stellatis induti vel sicut petioli
inflorescentia hypanthiaque pilis stipitato-dendroideus dense armati. Lamina 9.5—19 4.2—10.6 cm
elliptico-ovata 5(—7)-plinervata, supra glabra, subtus in venis secundariis tertiariisque pilis stellatis
modice puberuli. Panicula 4-8 cm longa multiflora; flores 5-meri; calycis tubus 0.5 mm altus, lobis
interioribus 1—1.5 mm longis late ovatis, dentibus exterioribus 1.5—2 mm eminentibus. Stamina
isomorphica glabra, thecis subulatis, poro dorsaliter inclinato; antherarum thecae 1 mm longae,
connectivum nec prolongatum nec appendiculatum. Ovarium 5-loculare et fere omnino inferum apice
in collo 0.25 mm alto modice pilis glanduliferis.

Shrub or small tree 2—6 m tall. Old branches terete, glabrous, and somewhat striate. Uppermost
cauline internodes, petioles, inflorescences, pedicels, and hypanthia densely covered with brown
penicillate-stellate and/or coarse dendritic hairs. Leaves of a pair equal to somewhat unequal in size;
petioles 0.7—1.5 cm long; blades chartaceous when dry, 9.5—19 x 4.2—10.6 cm, elliptic-ovate, apex
acuminate, base obtuse to broadly rounded, oblique and then slightly decurrent on the petioles, margin
undulate-denticulate varying to subentire, 5(—7)-plinerved, the innermost pair of elevated primaries
diverging from the median vein in subopposite or alternate fashion 0.5—1.6 cm above the blade base,
the transverse secondaries elevated and spaced 3-8 mm apart at the widest portion of the blade,
adaxially glabrous at maturity, abaxially beset with a copious cover of penicillate-stellate hairs on the
elevated primary veins and a moderate cover of stellate hairs on the prominulous network of trans-
verse secondary and higher order veins. Inflorescence terminal, 4-8 cm long, sometimes appearing
pseudolateral because of elongation of axillary shoots, paniculiform with ultimate branchlets termi-
nating in simple dichasia; bracts of rachis nodes paired, linear-oblong, 2-4 mm long, 0.25—1 mm
wide, essentially glabrous adaxially, copiously stellate-pubescent abaxially; bracteoles persistent,
typically 3 per pedicel, sessile, linear-oblong, 1-3 mm long, 0.25—0.75 mm wide, margin entire, gla-
brous adaxially and moderately stellate-pubescent abaxially. Pedicels obsolete or up to ca. 0.25 mm
long. Hypanthia (at anthesis) 22.5 mm long to the torus. Calyx tube 0.5 mm long, the calyx lobes
ovate to suborbicular, often bluntly mucronate at the apex, stellulate-puberulent on both surfaces,
1-1.5 x 1.5-2 mm; exterior calyx teeth 5, subulate, 1.5—2 mm long and conspicuously exceeding the
calyx lobes; torus glabrous on the adaxial face. Petals 5, glabrous, white, oblong-obovate, + rounded
apically, 3-4 mm long, 1.5—2 mm wide. Stamens 10, isomorphic, filaments glabrous, complanate,
1 mm long; anthers 1 mm long, 0.5 mm wide, yellow, laterally + compressed, bluntly subulate in dor-
sal and ventral views, + elliptic in profile view, + truncate apically with a dorsally inclined pore; con-
nective thickened dorsally but unappendaged. Ovary (at anthesis) completely inferior, 5-locular,
globose, apex fluted but becoming + rounded at maturity, apically crowned with an undulately lobed
collar 0.25 mm high that is minutely glandular-puberulent along the rim. Style straight, glabrous,
3.25 mm long, stigma capitellate. Berry globose, 4-5 mm in diameter. Seeds + triangular in outline,
angulate varying to somewhat rounded on the convex face, 0.5 mm long, white to tan, the testa
colliculose throughout, the lateral raphe extending the entire length of the seed.

PHENOLOGY. — The three known collections, made in January, February, and July, are in flower
and fruit.

DISTRIBUTION. — A little-collected rain forest species known from the southeastern corner of
Lim6n province in Costa Rica and the Nusagandi region of Comarca de San Blas province, Panama, at
150-350 m (Fig. 2).

PARATYPES. — PANAMA. Comarca de San Blas: Llano-Carti road, kilometer 16 along trail to
creek on the Caribbean drainage, 2 Feb. 1989, Almeda et al. 6514(CAS, MO, PMA, SCZ, US); head-
waters of Rio Nergala along continental divide, 11 Jan. 1985, de Nevers & Herrera 4514 (CAS, MO).

DISCUSSION. — Among described species of Miconia section Amblvarrhena, M. colliculosa is
most similar to a group of three allopatric species that includes M. calocoma Almeda of northeastern

35

ALMEDA: MICONIA

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bescence details at foliar base; C, flower (at anthesis) with one petal removed, * 10; D, flower as seen from above, = 5; E, petal
(adaxial surface), x 10; F, stamens, dorsal view (left), ventral view (middle), and profile view (right), x 19; G, mature berry, x 7;

H, seeds, x 40. (A-F from Almeda et al. 6154; G and H from the holotype.)

Volume 52, No. 4

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

36

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ALMEDA: MICONIA a7

Costa Rica, M. rupticalyx Wurdack of Venezuela, and M. wagneri J. F. Macbr. of Peru, Bolivia, and
northern Brazil (Almeda 1989a). All of these taxa differ from M. colliculosa in having 4-merous flow-
ers, an irregularly rupturing apiculate calyx, and other diagnostic reproductive characters. Miconia
calocoma, for example, has ventrally inclined anther pores and a 4-locular ovary. Miconia rupticalyx
and M. wagneri share the character state of dorsally inclined anther pores with M. colliculosa but both
have 2(—3)-locular ovaries. In addition, M. rupticalyx has reniform bracteoles whereas M. wagneri
has anther connectives that are dilated at the filament insertion into dorso-basal spurs.

ETYMOLOGY. — The epithet for this species is diminutive of the Latin word collinus, of a hill, in
reference to the little rounded or hillock-like elevations that make up the seed coat.

Miconia correae Almeda, sp. nov.
Fig. 3

TYPE. — PANAMA. Bocas del Toro: Trocha 3 de noviembre, near Paso de la Zorra shelter,
south and a bit west of Chiriqui Grande and ca. 2 km NE of the peak of Cerro Guayabo, 8°48'N,
82°14’-15’W,elev. ca. 1300 m, 5 Apr. 1978, Dressler 5806 (holotype: CAS!; isotype: PMA).

Section Jucunda. Frutex | m. Ramuli teretes sicut inflorescentia primum sparsiuscule vel modice
glandulis clavatis 1—1.75 mm longis induti demum glabrati. Folia in quoque pari disparilia; lamina
2.9-6.2 x 1.5—3.8 cm cordata vel ovata, 5—7-nervata, supra glabra, subtus pilis laevibus glanduliferis
sparsiusculi puberuli. Panicula 44.5 cm longa pauciflora; flores 4-meri; calycis tubus 0.5—1 mm
altus, lobis interioribus 1 x 1.5—2 mm triangularibus 1 mm altis, dentibus exterioribus subulatis
2.5-3 mm eminentibus. Stamina isomorphica glabra, thecis subulatis, poro dorsaliter inclinato;
antherarum thecae 2.5 mm longae, connectivo dorsaliter ad basim tuberculo 0.5 mm elevato ornato.
Ovarium 4-loculare et fere omnino inferum apice glabro.

Shrub to | m tall. The internodes terete, sparsely covered with spreading smooth glandular hairs
1 mm long when young like the inflorescence but becoming glabrous with age. Distal branchlet nodes
copiously setose with spreading hairs (glandular in part) 1—1.75 mm long. Leaves of a pair unequal in
size; petioles glabrous, |—2.9 cm long; blades membranaceous, somewhat brittle when dry, 2.9-6.2 =
1.5—3.8 cm, cordate varying to ovate, apex caudate-acuminate, base cordate to broadly rounded, mar-
gin ciliate-serrulate, 5—7-nerved, adaxially glabrous, abaxially sparingly and irregularly covered with
minute inconspicuous deciduous glandular hairs. Inflorescence terminal, 4—4.5 cm long, pani-
culiform, reportedly pendant with ultimate branchlets terminating in simple cymes or solitary flow-
ers; bracts of rachis nodes paired, ensiform to ligulate, 2—3 x 0.5 mm (including apical hair) essen-
tially glabrous on both surfaces; bracteoles narrowly lanceolate to subulate, 2 mm long (including
apical hair) and less than 0.5 mm wide, glabrous on both surfaces. Pedicels 1|—2 mm long at anthesis,
sparsely beset with spreading glandular hairs. Hypanthia (at anthesis) campanulate, 3-4 mm long to
the torus (vascular ring), glabrous throughout. Calyx lobes on flowering hypanthia 4, glabrous
throughout, rounded-deltoid, | mm long and 1.5—2 mm wide basally; exterior calyx teeth subulate,
2.5—3 mm long, adnate to and exceeding the calyx lobes. Petals 4, glabrous, white, elliptic-ovate,
rounded at the apex, 0.7—1 x 0.4-0.5 cm, the margin entire. Stamens 8, isomorphic; filaments gla-
brous, 2.5—3 mm long; anthers 2.5 mm long, yellow, subulate, rounded apically with a dorsally in-
clined pore; connective conspicuously thickened dorsally and prolonged basally into a caudiform
appendage 0.5 mm long. Ovary (at anthesis) completely inferior, 4-locular, oblong, glabrous at the
exposed apex. Style straight, glabrous, 7 mm long; stigma capitellate. Mature berry not seen.

PHENOLOGY. — The single known collection, which is in flower, was made in early April.

DISTRIBUTION. — The label on the type describes the habitat in western Panama as open rocky
areas at 1300 m (Fig. 2).

38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

FIGURE 3. Miconia correae Almeda. A, habit, x /2, with enlargement (right) showing nodal pubescence details; B, represen-
tative leaves (abaxial surfaces) from a node, * %4; C, portion of the inflorescence, ca. 2; D, petal, x 4; E, stamens, dorsal view
(left) and profile view (right), x ca. 11. (A—E from the holotype.)

ALMEDA: MICONIA 39

DISCUSSION. — The subulate anther thecae, dorso-basal connective prolongation, and greatly
developed calyx teeth that much exceed the calyx lobes dictate placement of M. correae in section
Jucunda as defined by Cogniaux (1891) and elaborated upon by Gleason (1958). Among described
species of Miconia, this new species most closely resembles M. zemurrayana Standl. & L. O. Wil-
liams of Mexico (Chiapas), Guatemala, El Salvador, and Honduras. They are similar in flower size,
petal color, and the elongate projecting calyx teeth. Miconia zemurrayana differs most conspicuously
in having glabrous internodes, narrower lanceolate leaves (0.5—1.7 cm) that are coarsely denticulate
(at least distally), 5S-merous flowers, unappendaged anther connectives, and ventrally inclined anther
pores. Another distinctive feature of M. zemurrayana is the presence of tufts of stipitate-stellate hairs
at the junction of the inner pair of elevated primaries with the median vein on the abaxial surface of
each leaf blade. In the protologue of M. zemurrayana, Standley and Williams (1950) gave no sectional
disposition for this species. Based on the characters enumerated above, it too can be assigned to sec-
tion Jucunda.

ETYMOLOGY. — This species is named for Mireya D. Correa, Curator of the herbaria at the Uni-
versity of Panama and the Smithsonian Tropical Research Institute. Professor Correa first brought the
only known collection of this species to my attention and has been a perennial source of assistance
during my field work in Panama over many years.

Miconia crocata Almeda, sp. nov.
Fig. 4

TyPE. — PANAMA. Cocle: Forested slopes above El Copé along abandoned road leading to the
continental divide, elev. 700-850 m, 23 Feb. 1988, Almeda et al. 5930 (holotype: CAS!; isotypes:
DUKE!, MEXU!, MO!, NY!, PMA!, TEX!, US!).

Section Cremanium. Frutex vel arbor parva 2—5 m. Ramuli quadrangulati demum teretes sicut
folia inflorescentia hypanthiaque plerumque glabri. Lamina 5—10 =< 1.4—3.4 cm anguste elliptica
trinervata. Panicula 1.5—2.5 cm longa multiflora; flores 5-meri; calycis tubus non evolutus, lobis
interioribus 0.5 mm longis rotundatis, dentibus exterioribus crassis appressis inframarginalibus.
Stamina isomorphica glabra poro ventraliter inclinato; antherarum thecae 0.5—0.25m
oblongo-cuneatae 4-porosae, connectivum vix (0.75 mm) prolongatum nec appendiculatum.
Ovarium 3-loculare omnino inferum apice costato sparsissime glanduloso-puberulo vel sparse
furfuraceo.

Shrub or small tree 2—5 m tall. Uppermost branchlet internodes glabrous, quadrate and conspicu-
ously carinate, becoming rounded and somewhat ridged in age. Leaves of a pair equal or slightly un-
equal in size; petioles glabrous, 0.6—3.2 cm long, blades membranaceous, glabrous on both surfaces,
5—10 x 1.4-3.4 cm, narrowly elliptic, apex acuminate, base narrowly acuminate, margin mostly en-
tire toward the blade base, otherwise obscurely crenulate, 3-nerved with an additional pair of incon-
spicuous inframarginal nerves evident above the blade base and becoming inconspicuous at the
acuminate apex, the transverse secondaries prominulous and spaced 1|—2 mm apart at the widest por-
tion of the blade. Inflorescence a terminal corymbiform panicle 1.5—2.5 cm long, the rachis glabrous,
quadrate and carinate; bracts and bracteoles evidently early deciduous and absent at anthesis but leav-
ing well-developed scars at upper nodes and at the base of floral pedicels. Pedicels 0.25 mm long or
not developed above the point of bracteole attachment, glabrous. Hypanthia (at anthesis) campanulate
to cupulate, 1.5 mm long to the torus (vascular ring), glabrous. Calyx lobes 5, glabrous throughout,
broadly rounded-undulate, 0.5 mm long and 0.5—0.75 mm wide basally, exterior teeth broadly deltoid
or evident as a thickening mostly less than 0.25 mm long, adnate to and mostly shorter than the calyx
lobes when dry. Petals 5, erect and concave, glabrous, yellow or yellow with a flush of red or maroon

40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

apically, oblong-ovate, rounded at the apex, 1.5 x 1—1.25 mm, the margin entire. Stamens 10, isomor-
phic, incurved toward the central axis of the flower; filaments glabrous, complanate, tapered from
base to apex, saffron yellow when dry, 2 mm long; anthers 4-celled, 0.5 mm long, 0.25 mm wide at
the apex, yellow or yellow flushed with red apically, + infundibuliform to obliquely cuneate in profile
view, the pore + quadrate and strongly inclined ventrally; connective thickened and prolonged
0.75 mm below the anther thecae but lacking appendages. Ovary (at anthesis) inferior, 3-locular,
globose, apex fluted and sparsely glandular- or furfurate-puberulent but becoming rounded to trun-
cate and glabrate in fruit. Style straight, glabrous, 2 mm long; stigma truncate to capitellate. Berry
globose, blue-green becoming white with a faint flush of blue when mature, 4-5 mm in diameter.
Seeds + pyriform, 0.5 mm long, beige, tuberculate.

PHENOLOGY. — Flowering sporadically from January through September; fruiting specimens
have been collected from January through May and in October.

DISTRIBUTION. — Low montane cloud forests along the continental divide in Coclé province
from the El Copé region northeast to Los Volteaderos at 480—1000 m (Fig. 5).

PARATYPES. — PANAMA. Cocleé: Slopes above El Copé along abandoned road leading to the
continental divide, 8°38'N, 80°38'W, 24 Jan. 1989, Almeda et al. 6388 (CAS, DUKE, MICH, MO,
NY, PMA, US); about 7—-10.5 km beyond El Copé in Omar Torrijos National Park along rocky trail to
Rio Blanco and Limon beyond Alto Calvario, 21 Feb. 1996, Almeda et al. 7649 (CAS, MO, PMA,
US); 12.4 km N of La Pintada on the road from Penonomé to Coclecito on trail through remnant forest
on the continental divide at Los Volteaderos, 23 Feb. 1996, Al/meda et al. 7680 (CAS, PMA); El Copé,
along gravel road to the right before the sawmill, 18 Oct. 1979, Antonio 2204 (CAS, MO); region del
Copé, 24 May 1981, Correa et al. 4277 (CAS, PMA); Alto Calvario region, 4.5 miles N of El Cope,
8°38’/N, 80°36'W, 12 Sep. 1987, Croat 67517 (CAS, MO); Alto Calvario about 6 km N of El Copé,
8°39’/N, 80°36'W, 23 June 1988, Croat 68828 (CAS, MO); along Atlantic side of continental divide
above El Copé, 8°40’N, 80°37’W, 25 Jul. 1983, Miller et al. S35 (CAS, MO); Atlantic slope of the
continental divide above El Copé, 8°40'N, 80° 36’W, 13 Feb. 1982, Knapp & Dressler 3407 (CAS,
MO); El Copé, Division continental arriba de Barrigon y el aserradero viejo, 27 Apr. 1992, Pena etal.
369 (CAS, PMA); above EI Potroso sawmill N of El Copé, 13 May 1981, Sytsma & Andersson 4561
(CAS, PMA); 4.5 miles N of El Copé near the old sawmill, 8° 38’N, 80°35'W, 8 Apr. 1988, Thompson
4775 (CAS, CM).

DISCUSSION. — Among the species of section Cremanium characterized by a 3-locular ovary
and obovoid, 4-celled anthers that open by a wide terminal pore, M. crocata appears to be most similar
to M. chiriquiensis Almeda of Costa Rica and Panama and M. rubens (Sw.) Naudin of Jamaica, Co-
lombia, and Venezuela. The latter differs from M. crocata by the somewhat swollen cauline nodes,
ferrugineous pubescence on distal nodes and juvenile foliage, bracteoles with fimbriate margins,
dioecious floral condition, and glabrous ovary apex. Miconia crocata seems closest to M.
chiriquiensis. They share similarities in anther and stigma morphology, posture and orientation of
petals and androecium at anthesis. In both species the petals are erect and concave and the filaments
are geniculate in a way that closely juxtaposes the anther pores in a ring around the stigma. However,
M. chiriquiensis has leaf blades that are ciliate-serrulate distally, caudate-acuminate apically, and
furfurate-punctate abaxially. Miconia chiriquiensis also differs in a number of other diagnostic char-
acters. It has an openly branched elongate paniculate inflorescence, suborbicular petals, a glabrous
ovary apex, and seeds that superficially appear to have a smooth testa but are, in fact, minutely
papillate. Miconia crocata and M. chiriquiensis are allopatric and have different elevational ranges.
The latter ranges from the Cordillera de Talamanca in Costa Rica southeast to the Volcan Bart region
of western Panama and consistently occurs at higher elevations (1600-2500 m).

All Panamanian and some Costa Rican populations of M. chiriquiensis also have white petals and
anthers, but some recent Costa Rican collections (Almeda & Anderson 5322; Grayum 10333) have
petals and stamens that are bright yellow like those of M. crocata.

ALMEDA: MICONIA

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cence showing a fully opened flower, x ca. 8; D, petals (adaxial surface), x ca. 22; E, stamens, profile view (left), ventral view
(center), and dorsal view (right), x 24; F, mature berry, x 6; G, seeds, x 40. (A—G from the holotype.)

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

42

Volume 52, No. 4

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ALMEDA: MICONIA 43

ETYMOLOGY. — Because yellow is such an uncommon petal color in the tribe Miconieae, the
name chosen for this new species emphasizes this feature. The epithet crocata is derived from the
Greek word, krokotos, saffron-yellow, in reference to the striking color of the corolla and androecium,
especially upon drying.

Miconia jefensis Almeda, sp. nov.
Fig. 6

TYPE. — PANAMA. Panama: Cerro Jefe, along summit road and along trail into the Chagres
Valley, ca. 900 m, 19 Feb. 1988, Almeda et al. 5826 (holotype: CAS!; isotypes: MO!, NY!, PMA!).

Section Miconia. Frutex 2-4 m. Ramuli primum paulo compressi demum sulcato-quadrangulati
sicut petioli foliorum subtus venae primariae inflorescentiaque dense pilis penicillato-stellulatis
induti. Lamina 13.5—41 x 7-14 cm elliptica vel elliptico-ovata 5—7-plinervata supra glabra, subtus in
venis secundariis tertiariisque pilis stellulatis modice puberuli. Panicula 6—14 cm longa multiflora;
flores 5-meri; calycis tubus 0.25 mm altus, lobis interioribus 0.25 longis late rotundatis, dentibus
exterioribus crassis adhaerentibus lobos interiores aequantibus. Stamina isomorphica glabra, thecis
subulatis, poro paulo dorsaliter inclinato; connectivum nec prolongatum nec appendiculatum.
Ovarium 5-loculare et 4/5 inferum apice sparsicuscule glanduloso-puberulo.

Shrub 2-4 mtall. Uppermost branches compressed-rounded becoming bisulcate to rounded-
quadrate with age, the young branchlets, petioles, elevated primaries of abaxial foliar surfaces, and in-
florescence rachis and pedicels densely covered with a brown scurfy indument of
penicillate-stellulate hairs. Leaves of a pair somewhat unequal in size; petioles |—4(—7) cm long,
blades subcoriaceous when dry, 13.5-41 « 7-14 cm, elliptic to elliptic-ovate, apex long acuminate to
attenuate, base varying from subcordate to tapering and abruptly rounded at the petiole junction, mar-
gin entire, 5—7-plinerved, the innermost pair of elevated primaries diverging from the median vein in
alternate fashion (2—)4.2—-8.5 cm above the blade base, the transverse secondaries elevated and spaced
5—8 mm apart at the widest portion of the blade, adaxially glabrous at maturity, abaxially moderately
covered with a brown scurfy-stellulate indument on the secondary and higher order veins. Inflores-
cence a terminal multiflowered panicle 6—14 cm long, divaricately branched at the node initiating the
inflorescence; bracts of the rachis nodes persistent, 1.54.5 mm long, 0.25—1 mm wide, glabrous
adaxially, glabrous or sparsely covered with scurfy puberulence abaxially; bracteoles sessile and per-
sistent, paired, trimerous, or quaternate, sometimes fused basally to form a shallow nodal collar,
subulate, 1-1.25 mm long, 0.5 mm wide basally, essentially glabrous adaxially and sparingly
stelluate-furfuraceous to glabrous abaxially. Pedicels nearly obsolete or up to 1 mm long. Hypanthia
(at anthesis) 1—-1.5 mm long to the torus, copiously to moderately stellulate-furfuraceous. Calyx tube
0.25 mm long, the calyx lobes rounded-triangular, 0.25 mm long; exterior calyx teeth 5, bluntly
subulate, up to 0.25 mm long, equaling or somewhat shorter than the calyx lobes but typically ob-
scured by the dense indument; torus fimbrillate-puberulent or glandular-puberulent. Petals 5, gla-
brous, magenta, elliptic-ovate to oblong, rounded apically, 2-3 mm long, 0.75—1 mm wide. Stamens
10, isomorphic, filaments glabrous, complanate, constricted distally just below the anther thecae,
0.75—1 mm long; anthers | mm long, 0.25 mm wide, pale yellow, linear-oblong, rounded to truncate
at the apex with a somewhat dorsally inclined terminal pore; connective thickened dorsally but
unappendaged. Ovary (at anthesis) 4/5 inferior, 5-locular, globose, apex deeply fluted but becoming
rounded at maturity, densely white-papillate with a few brown glandular hairs. Style typically de-
clined to one side of the flower, glabrous, 2.5—3 mm long; stigma capitellate. Berry pink but turning
blue-purple when mature, 3-4 mm long and 3-4 mm in diameter. Seeds angular-pyramidate, 0.5 mm
long, brown or tan, smooth with finely verruculose angles on the convex face.

44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

PHENOLOGY. — Flowering sporadically from February through July; fruiting specimens have
been collected in February, July, and September.

DISTRIBUTION. — Uncommon in low montane cloud forests in the Cerro Jefe and Cerro Azul re-
gions of Panama province at 700—1,000 m (Fig. 2).

PARATYPES. — PANAMA. Panama: About 29-30 km beyond the Interamerican Highway off of
a dirt road on Altos de Pacora, 26 Feb. 1996, Almeda et al. 7696 (CAS, MO, PMA); Vicinity of Cerro
Jefe, along road to summit, 9°14’N, 79°22’W, 8 Jul. 1987, Croat 67079 (CAS); 3—3.5 miles NE of
Altos de Pacora and 11.1—11.6 miles beyond Lago Cerro Azul, 19 June 1988, Croat 68627 (CAS,
MO); Sendero “El Cantar,” borde de quebrada, Cerro Azul, Parque Nacional Chagres, 26 Jul. 1991,
Carrasquilla et al. 3267 (CAS, PMA); region de Cerro Jefe, area cercana la antena, 13 Jul. 1994,
Galdames & Montenegro 1389 (CAS, SCZ); region de Cerro Jefe, area cercana al limite con la
Urbanizacion Altos de Cerro Azul, 13 Jul. 1994, Galdames & Montenegro 1409 (CAS, SCZ);
headwaters of the Rio Utivé, Cerro Jefe, 2 km from last branch in road to summit, 13 Sep. 1981,
Knapp 1209 (CAS, MO).

DISCUSSION. — Miconia jefensis is related to a group of species that includes M. iteophylla
Almeda, M. ligulata Almeda, and M. peltata Almeda. Like M. jefensis, two of these taxa are restricted
to Panama; M. ligulata ranges from Nicaragua to Venezuela (Almeda 1989b). All of these species
share a scurfy puberulent indument, linear or oblong petals, unappendaged anthers, a 5-locular ovary,
a torus that is puberulent adaxially, and seeds that are nearly identical in size and shape. In overall as-
pect, M. jefensis is most like M. ligulata. These two species have angular-pyramidate seeds like the
other members of this alliance but they alone share the fine verruculose ornamentation at the angles of
the seed testa.

I was inclined to regard initial collections of M. jefensis as extraordinary variants of M. ligulata.
Study of additional material, and the opportunity to examine more than one population of this entity in
the field, revealed significant differences in a number of characters that are consistent with the kinds
of characters that distinguish closely related species in other complexes within the genus. Miconia
jefensis is a coarse, robust species with stout nodes that measure 6—10 mm on the widest face. Other
diagnostic characters include its magenta petals, 5—7-plinerved leaves with a base varying from
subcordate to abruptly rounded at the petiole junction, an inflorescence that is divaricately branched at
the node initiating the inflorescence, and an ovary apex that is persistently white-papillate intermixed
with a few brown glandular hairs at maturity. Plants of M. /igu/ata, in contrast, are slender in aspect
with thinner cauline nodes that measure 3—5 mm at the widest face. This species has white petals,
5-plinerved leaves with a base that is gradually tapered and decurrent on the petiole, an inflorescence
that typically branches 1.5—3.5 cm above the initiating node, and an ovary apex that is glabrous in
fruit.

Miconia jefensis appears to be endemic to the low montane cloud forest that covers the volcanic
region of Cerro Jefe and Cerro Azul, Panama. Hence, this narrowly endemic species joins a varied as-
semblage of flowering plants restricted to this region (Lewis 1971), including several recently de-
scribed species of Melastomataceae such as Adelobotrys jefensis Almeda (Almeda 1981), Miconia
morii Almeda (described herein), Miconia peltata Almeda (Almeda 1989b), Tessmannianthus
carinatus Almeda (Almeda 1989c), and Topobea hexandra Almeda (Almeda 1990).

ETYMOLOGY. — The specific epithet is derived from the type locality, Cerro Jefe, where most of
the collections of this species have been made.

ALMEDA: MICONIA 45

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FiGurE 6. Miconia jefensis Almeda. A, habit, x 1/3; B, enlargement of detail on cauline internode; C, representative leaf
(abaxial surface), x %; D, fully opened flower, x ca. 13; E, petal (adaxial surface), x ca. 23; F, stamens, profile view (left) and
dorsal view (right), x 18; G, mature berry, x 7; H, seeds, x 40. (A—F from the holotype; G and H from Almeda et al. 7696.)

46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

Miconia morii Almeda, sp. nov.
Fig. 7

TYPE.—PANAMA. Panama: Cerro Jefe, along trail on ridge running NE from summit, cloud for-
est dominated by Clusia spp. and Colpothrinax cookii, ca. 1000 m, 18 Dec. 1974, Mori et al. 3773
(holotype: CAS!; isotypes: MO, WIS!).

Section Tamonea. Frutex 1.2-3m. Ramuli obtuse tetragoni demum teretes sicut petioli
inflorescentia hypanthiaque modice vel dense pilis stellulatis induti. Lamina 6.5—10.5 « 2.5-5 cm
elliptica vel elliptico-ovata trinervata, supra glabra, subtus in venulis superficieque sparsiuscule
caduceque stellato-puberulo. Panicula 8-11 cm longa multiflora; flores 5-meri; petala extus pilis
stellulatis modice induta; calyx 2—2.5 mm longus truncatus, dentibus exterioribus obscuris (0.25 mm)
omnino adhaerentibus non eminentibus. Stamina isomorphica, thecis subulatis, poro ventraliter
inclinato; antherarum thecae 44.25 mm longae, connectivum dorsaliter ad basim paulo elevatum
ecalcaratum. Ovarium 5-loculare et 2 inferum apice in collo 0.25 mm alto sparsissime glanduloso
puberulo.

Shrub 1.2—3 m tall. Uppermost cauline internodes compressed-rounded to obtusely quadrangu-
lar becoming terete with age, the uppermost branchlets, vegetative buds, young leaves and petioles,
inflorescence branches, hypanthia, and pedicels moderately to densely appressed-puberulous with
brown stellulate hairs 0.7—1 mm in diameter. Leaves ofa pair equal to somewhat unequal in size, peti-
oles 0.9—1.5 cm long, blades subcoriaceous when dry, 6.5—10.5 x 2.5—5 cm, elliptic to elliptic-ovate,
apex acuminate, base obtuse to rounded, margin entire, adaxially glabrous or essentially so, abaxially
moderately to sparsely stellulate-puberulous but sometimes becoming glabrate with age, 3-nerved,
the transverse secondary veins elevated and spaced 4.5—8 mm apart at the widest portion of the blade.
Inflorescence a terminal multiflowered panicle 8-11 cm long, branching at or 2—2.5 cm above the
node initiating the inflorescence; bracts and bracteoles evidently early deciduous and not seen. Pedi-
cels 0.5 mm long. Hypanthia (in fruit) 4-5 mm long to the torus, calyx tube 2—2.5 mm long, flangelike
apically and sometimes splitting vertically down toward the torus at one or more points, the calyx
lobes essentially obsolete, only evident as a truncate rim; exterior calyx teeth 5, barely evident as blunt
callosities ca. 0.25 mm long; torus essentially glabrous adaxially. Petals 5, adaxially glabrous,
abaxially densely stellulate-puberulous, oblong-obovate, cucullate distally and rounded at the apex,
5.5—7 mm long, 4—5 mm wide distally. Stamens 10, isomorphic, filaments glandular-puberulent ba-
sally, 3 mm long; anthers 44.25 mm long, ca. 0.75 mm wide at the base, linear-oblong to subulate,
rounded at the apex and opening by a ventrally inclined pore; connective elevated dorso-basally into
an elongate padlike thickening | mm long. Ovary (in fruit) /2-inferior, 5-locular, subglobose, apex
conical, inconspicuously glandular-puberulent, terminating in a shallowly unduiate collar 0.25 mm
high that is typically glabrate but sometimes adorned with a few minute glandular hairs on the rim.
Style straight, glandular-puberulent along the basal third of its length, 6 mm long, stigma capitate.
Berry subglobose, 4—6 mm long, 5—7 mm in diameter. Seeds + triangular in outline, rounded to
angulate on the convex face, 0.5—1 mm long, brown, smooth with a dull luster, the lateral raphe ex-
tending the entire length of the seed.

PHENOLOGY. — The only flowering collection was made in March; fruiting specimens have
been collected from December through May.

DISTRIBUTION. — Known only from the summit and vicinity of Cerro Jefe just east of the Canal
Area in Central Panama at 900—1,000 m (Fig. 5).

PARATYPES. — PANAMA. Panama: Vicinity of Cerro Jefe, near tower, 23 May 1980, Antonio
4725 (CAS, MO); Cerro Jefe, 28 Sep. 1986, Aranda 154(PMA, US); road from summit of Cerro Jefe,
9°14'N, 79°23'W, 20 Jan. 1984, Churchill 4298 (CAS, MO); Cerro Jefe, by radio tower, 15 Mar.

ALMEDA: MICONIA 47

2
3
2
z
&
i
te
=

FIGURE 7. Miconia morii Almeda. A, habit, x 1/3; B, representative leaf (abaxial surface), x 1; C, petal (adaxial surface), x 6;
D, stamens, profile view (left) and dorsal view (right), x 14; E, mature berry as seen from above, x 5; F, mature berries in profile
view showing flangelike calyx (right) that sometimes splits in an irregular vertical fashion (left), x 4; G, seeds, x 20. (A from the

holotype and Churchill 4298; B from the holotype; C and D from Folsom et al. 2525; E and F from Churchill 4298; G from
D’Arcy and McPherson 16060.)

48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

1985, D’Arcy & McPherson 16060 (CAS, MO); top of Cerro Jefe, 23 km N of Pan-American Hwy.,
11 Apr. 1977, Folsom et al. 2325 (CAS, MO).

DISCUSSION. — Among the species of section Tamonea, M. morii is morphologically most simi-
lar to M. acuminifera Triana of western Colombia (Depts. of Caldas south to Narifio). These two spe-
cies have similar leaf shape, hypanthial and petal indument, calyx details, and seed morphology. They
are also similar in having isomerous flowers (ovary 5-locular and petals 5 in number). In M.
acuminifera, however, the flowers are pleiostemonous (vs. diplostemonous in M. morii) and the sta-
mens are dimorphic in size (vs. isomorphic in M. morii). Other differences that distinguish these two
species involve floral or fruit characters. In M. acuminifera the style and filaments are glabrous, the
toral vascular ring to which stamens and petals are attached is glandular-puberulent adaxially, and the
connective lacks the elevated padlike thickening so typical of M. morii (Fig. 5D). The ovary summit in
both M. morii and M. acuminifera is glandular-puberulent, although inconspicuously so at times, but
the latter lacks the well-developed collar that surrounds the style base of the former.

There are other modal, but more subtle, differences that can be used to separate these two species.
In M. acuminifera the leaf blades are typically larger (14—28 x 5—12.5 cm), the fruiting hypanthia are
strongly constricted distally, and the stellate indument on flowering hypanthia is so dense that it con-
ceals the actual surface. The leaf blades of M. morii are 6.5—10.5 x 2.5—5 cm, the fruiting hypanthia
are tapered from the base to the apex (Fig. SF), and the hypanthial indument does not conceal the ac-
tual surface. In addition to being allopatric, M. acuminifera appears to grow at higher elevations. Only
one of the collections examined was gathered at 950 m; all others were collected at 1,550—2,100 m.

ETYMOLOGY. — This species is named for Scott A. Mori (1941— ) collector of the type of this
species and other interesting Melastomataceae during his early years of botanical field work in
Panama.

Miconia talamancensis Almeda, sp. nov.
Fig. 8

TYPE. — COSTA RICA. Cartago/San José border: Cordillera de Talamanca, Villa Mills in the
vicinity of La Georgina, ca. 9750 ft (2972 m), 6 Mar. 1981, Almeda & Nakai 4777 (holotype: CAS!;
isotypes: CR!, INB!, MO!).

Section Cremanium. Arbor 6-15 m. Ramuli primum obtuse sulcato-quadrangulati demum
teretes sicut petioli folia novella inflorescentia hypanthiaque pilis asperis dense vel modice induti
demum glabrati. Lamina 4.5—12 x 1.9-3.8 cm oblongo-ovata 5-nervata, supra glabra, subtus in
venarum primariarum axillis (et supra axillis) modice setosa pilis simplicibus 1—3 mm longis
persistentibus. Panicula 6—9.5 cm multiflora; flores 5-meri; calycis tubus 0.25 mm altus, lobis
interioribus 0.5 longis, dentibus exterioribus crassis inframarginalibus. Stamina in dimensionibus
subisomorphica glabra, antherarum thecae 1.5—1.75 mm oblongae latae 4-porosae, connectivo
prolongato dorsaliter inconspicue hebeti-dentato. Ovarium 3-loculare et ’2 inferum apice glabro.

Tree 6-15 m tall. Uppermost cauline internodes quadrate becoming rounded-quadrate with age,
the uppermost branchlets, vegetative buds, young leaves and petioles, inflorescence branches, and
bracteoles densely covered with a rusty brown indument of asperous-headed, dendritic hairs. Leaves
of a pair equal or slightly unequal in size, petioles 1.14.1 cm long, blades coriaceous when dry,
4.5—-12 x 1.9-3.8 cm, oblong-ovate, apex caudate-acuminate to attenuate, base rounded, margin
callose-serrulate, adaxially glabrous and somewhat bullate-reticulate, abaxially scattered
asperous-lepidote on the actual surface but copiously covered with rusty brown dendritic or
asperous-headed hairs on the elevated primary veins and moderately covered with simple spreading
hairs (1-3 mm long) where the primary veins diverge at the blade base and commonly extending for

ALMEDA: MICONIA 49

B
Y awe
Paper,

[YH Su
ke ey.
ao” eR

FIGURE 8. Miconia talamancensis Almeda. A, habit, x 1; B, representative leaf (abaxial surface), x 1; C, enlargement show-
ing pubescence detail at a leaf base (abaxial surface); D, ultimate dichasial inflorescence unit showing flower buds, 6; E, petal
(adaxial surface), x 9; F, antepetalous stamen, ventral view (left) and profile view (right), x ca. 11; G, antesepalous stamen, ven-
tral view (left) and profile view (right), x ca. 11; H, seeds, x ca. 18. (A—H from the holotype.)

50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

some distance up the blade along the primaries (especially the median vein), 5-nerved (the outermost
pair often inconspicuous), the transverse secondary veins elevated and spaced 1.54 mm apart at the
widest portion of the blade. Inflorescence a terminal multiflowered panicle 6—9.5 cm long, branching
1.5-2.5 cmabove the node initiating the inflorescence; bracts of the rachis nodes early deciduous and
not seen; bracteoles also early deciduous and rarely seen at anthesis, paired, elliptic-oblong to ob-
long-obovate, 0.75—1.5 mm long, 0.25—0.5 mm wide (at the widest point). Pedicels 0.5—1.25 mm
long. Hypanthia (at anthesis) 2—2.5 mm long to the torus, moderately covered with rusty brown den-
dritic or asperous-headed hairs that typically do not completely conceal the actual surface. Calyx tube
0.25 mm long, the calyx lobes deltoid to broadly rounded-deltoid, hyaline, 0.5 mm long; exterior ca-
lyx teeth 5, bluntly triangular, 0.25 mm long, shorter than and not obscuring the calyx lobes; torus gla-
brous adaxially. Petals 5, glabrous, white, obovate to + suborbicular, rounded and often retuse or
emarginate apically, 1.5-2 mm long, 1—1.5 mm wide. Stamens 10, subisomorphic, filaments gla-
brous, geniculate above the middle, constricted distally about 2/3 of the way up from the base,
2—2.5 mm long; anthers 1.5—1.75 mm long, 0.75 mm wide, white, oblong-obovate, widest above the
middle, 4-celled, rounded to subtruncate at the apex and opening by a wide ventrally inclined opening;
connective lobulate-thickened dorso-basally and prolonged ventrally below the thecae into two lat-
eral lobes 0.24 mm long. Ovary (at anthesis) 2-inferior, 3-locular, globose, apex + rounded and gla-
brous at maturity. Style straight, glabrous, 3-3.5 mm long; stigma capitellate. Berry globose,
greenish-white becoming blue-black when mature, 5—6 mm in diameter. Seeds ovoid to ellipsoid,
0.5—0.75 mm long, tan, smooth with a dull luster, the lateral raphe extending for much of the seed
length.

PHENOLOGY. — Collected in flower from February through July; in fruit during the months of
March, June, and September.

DISTRIBUTION. — Locally common in high montane cloud forests and in rocky areas bordering
paramo from near Villa Mills to Cerro Kamuk on the Cordillera de Talamanca of Costa Rica southeast
to the Volcan Bart region of western Panama at 2900-3350 m (Fig. 5).

PARATYPES. — COSTA RICA. Cartago/San José border: About 3 km W of Villa Mills, 10 Mar.
1981, Almeda & Nakai 4832 (CAS, CR); Cordillera de Talamanca, La Georgina, 27 Jul. 1972, Kesel
& Sauer 5307 (CAS); Limon: Cordillera de Talamanca, SW foot of Cerro Kamuk, 9°16'N,
83°02'30"W, 24 Mar. 1984, Davidse et al. 25970 (CAS); Chirrip6 National Park, N end of Loma
Larga, 15 Feb. 1983, Garwood et al. 1262 (BM, MO). Puntarenas: Cordillera de Talamanca, across
the Panamerican Hwy. from La Georgina, 23 Feb. 1991, Almeda etal. 6785 (CAS, CR, MEXU, MO).
San José: Along Interamerican Hwy. ca. 25 km SW of road to La Cima and 4.1 km NW of Cerro
Asuncion, 9°36'N, 83°46’'W, 11 Sep. 1979, Stevens 14269 (CAS). PANAMA. Chiriqui: 12 miles
above Boquete on road to Volcan Baru, 18 May 1976, Croat 34886 (CAS); La Nivera, below summit
of El Bart, 14 Mar. 1979, D’Arcy & Hammel 12467 (CAS); top of high ridge N of summit of Volcan
Cerro Pavon, 15 Mar. 1979, Hammel & D’Arcy 6436 (CAS); Volcan Baru, along road to summit,
8°45’'N, 82°30'W, 10 June 1986, McPherson 9487 (CAS).

DISCUSSION. — Miconia talamancensis has white, obovate to suborbicular petals; white,
4-celled anther thecae; ovoid to ellipsoid seeds with a dull luster; and oblong-ovate leaves with a con-
centration of simple spreading hairs where the primary veins diverge from one another at the abaxial
base of the blade. In all of these characters, M. talamancensis is very similar to M. schnellii Wurdack,
and there is no doubt that they are closely related. These two species differ in features of the indument,
exterior calyx teeth, and anther details. In M. talamancensis the uppermost internodes, young leaves,
petioles, and inflorescences are densely covered with a rusty brown indument of asperous-headed or
dendritic hairs that are most reminiscent of the dendritic hairs with short arms illustrated by Wurdack
(1986:64). Miconia talamancensis also has exterior calyx teeth that do not exceed or obscure the calyx
lobes, and the septum separating the four anther cells does not conspicuously protrude beyond the an-
ther apex. In M. schnellii the upper internodes and inflorescence branches are completely glabrous,

ALMEDA: MICONIA 51

the calyx teeth exceed and obscure the calyx lobes, and the septum of the anther thecae protrudes be-
yond the anther apex.

In most collections of M. talamancensis the simple hairs at the abaxial leaf base commonly ex-
tend for some distance up the blade along the median vein. I have never observed this kind of pubes-
cence distribution in any population or herbarium collection of M. schneillii. | have found both of these
species growing side by side on the Cordillera de Talamanca in Costa Rica with no breakdown or
intermediacy in the character differences noted above.

Although M. talamancensis and M. schnellii have overlapping geographic and elevational
ranges, there are some modal differences worthy of note. The latter grows on Costa Rica’s Cordillera
de Talamanca and on the slopes of Volcan Irazu and Volcan Turrialba at elevations of 1,980—3,200 m.
In Costa Rica, M. talamancensis occurs only on the Cordillera de Talamanca with populations extend-
ing southeast to the Volcan Baru region of Panama. All collections of this species have been made at
2,900-3,350 m.

ETYMOLOGY. — The specific epithet is derived from Cordillera de Talamanca, a plutonic uplift
dominating the mountainous backbone of Costa Rica, where many of the collections of this species
have been made.

Miconia vestita Almeda, sp. nov.
Fig. 9

TYPE.—COSTA RICA. San José: Ridgetop due E of Finca Chacon near San Gerardo de Dota,
elev. 2500 m, 9 Mar. 1995, Almeda 7399 (holotype: CAS!; isotype: CR!).

Section Cremanium. Arbor parva 2—S m. Ramuli teretes sicut petioli foliorum venae primariae
subtus inflorescentiaque pilis stipitato-stellatis erectis dense vel modice armati et pilis laevibus
glanduliferis sparse intermixtis. Lamina 8.7—14.4 x 4-6.9 cm elliptico-ovata 5—7-nervata, supra
glabra, subtus in venis secundariis tertiariisque pilis stipitato-stellatis modice puberuli. Panicula
9—12 cm longa multiflora; flores 5-meri; calycis tubus 0.25 mm latus, lobis interioribus 0.5—1 mm
longis oblongo-ovatis, dentibus exterioribus minutis inconspicuis inframarginalibus. Stamina
isomorphica glabra, antherarum thecae 1I—1.5 x 0.25 mm oblongae poro ventraliter inclinato,
connectivo dorsaliter ca. 0.25 mm bilobulato. Ovarium 3-loculare et 2/3 inferum apicem versus
sparsiuscule glandulis vel paullulo stellatis armatum.

Small openly branched tree 2—5 m tall. Uppermost cauline internodes and inflorescence rachis
copiously covered with a rusty brown indument of dendritic and penicillate-stellate hairs (0.5 mm
long) intermixed with smooth spreading glandular hairs (mostly 1.5 mm long). Leaves of a pair
mostly equal in size; petioles 2.5—3.3 cm long; blades subcoriaceous when dry, 8.7—14.4 x 46.9 cm,
elliptic to elliptic-ovate, apex acuminate to cuspidate, base obtuse to rounded, margin inconspicu-
ously serrulate, 5—7-nerved, the transverse secondaries elevated and spaced 24 mm apart at the wid-
est portion of the blade, adaxially glabrous or sparsely and irregularly beset with dendritic or
stipitate-stellate hairs in the channels created by the impressed primary veins, abaxially beset with a
dense to moderate cover of rusty colored dendritic and penicillate-stellate hairs on the elevated pri-
mary veins and on the prominulous network of transverse secondary and higher order veins. Inflores-
cence terminal, 9-12 cm long, paniculiform with ultimate branchlets terminating in simple dichasia;
bracts of rachis nodes paired judging from nodal scars but evidently early deciduous and not seen,
bracteoles early- or tardily-deciduous, oblong to narrowly oblanceolate, 2—2.5-3 mm long, 0.5 mm
wide, glabrous adaxially, moderately to copiously stipitate-stellulate abaxially. Pedicels
0.25—0.5 mm long. Hypanthia (at anthesis) 2.5—-3 mm long to the torus, moderately to sparsely cov-
ered with stellate and stipitate-stellate hairs that are sometimes intermixed with a few spreading glan-

52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

FIGURE 9. Miconia vestita Almeda. A, habit, ca. 4; B, representative leaf (abaxial surface), with enlargement of pubes-
cence detail, x 1; C, open flowers showing posture of petals and stamens, * ca. 4; D, hypanthium in profile view with petals and
androecium removed, * ca. 7; E, petal, x 10; F, stamens, ventral view (left) and partial profile view (right), x ca. 17; G, berry as
seen from above, * ca. 6; H, séeds, x ca. 10. (A—-H from the holotype.)

ALMEDA: MICONIA 53

dular hairs. Calyx tube 0.25 mm long, the calyx lobes oblong-ovate, rounded at the apex, abaxially
glabrous, adaxially glabrate or sparingly stellate-pubescent at the base and adjacent to the calyx teeth,
0.5—1 x 0.5—0.75 mm; exterior teeth 5, subulate to triangular, 0.5 mm long, typically shorter than the
calyx lobes but often obscured by an indument of stipitate-stellate and glandular hairs; torus glabrous
on the adaxial face. Petals 5, glabrous, white, reflexed, obovate, rounded and emarginate apically,
2-2.5 mm long, 2 mm wide. Stamens 10, isomorphic, reflexed, forming a wide circle around the style
and lying close to the expanded petals, filaments geniculate, glabrous, complanate, 2 mm long; an-
thers |—-1.5 mm long, 0.25 mm wide, white, linear-oblong, + truncate apically with a + ventrally in-
clined pore; connective thickened and elevated dorso-basally into a bilobed appendage mostly less
than 0.25 mm long and prolonged ventro-basally into a caudiform lobe 0.25 mm long at the base of
each anther sac. Ovary (in fruit) 2/3 inferior, 3-locular, globose, apex + rounded and sparingly beset
with a few stellate and/or glandular hairs. Style straight, glandular-puberulent, 3.5 mm long, stigma
capitate. Berry globose, 3.54 mm in diameter. Seeds irregularly angulate-pyramidate, 1 mm long,
the testa smooth and + vernicose, the lateral raphe extending the entire length of the seed.

PHENOLOGY. — The type, which was collected in early March, is in flower and young fruit.

DISTRIBUTION. — Known only from the type locality on a ridgetop dominated by Quercus
bumelioides Liebm. [Q. copeyensis C. H. Muller] near San Gerardo de Dota on the Cordillera de
Talamanca, Costa Rica at 2500 m (Fig. 2).

DISCUSSION. — Although M. vestita is a typical member of section Cremanium in having oblong
2-celled anthers and permanently geniculate filaments above the middle, it does not appear to be par-
ticularly close to any of the 209 species currently placed in this section. In foliar size and shape there is
a superficial resemblance to M. acanthocoryne Wurdack of Colombia, but that species has a
rubiginose-furfuraceous indument on upper branches, petioles, and inflorescences, 4-celled anthers
with broad pores and 2-locular ovaries.

The most striking feature of M. vestita is the indument on the abaxial foliar surface which consists
of rusty brown dendritic or penicillate-stellate hairs reminiscent of a miniature forest of trees. The an-
thers are also noteworthy because the thecae appear thin-walled, somewhat collapsed but not mal-
formed, and largely devoid of pollen. It is possible that all the flowers on the two individuals that I
encountered in the field had been relieved of their pollen by buzzing bees, but it is also possible that I
collected functionally pistillate individuals of a dioecious species. Non-functional stamens are pro-
duced on the pistillate flowers of other dioecious species of Miconia, all of which belong to section
Cremanium and occur at high elevations. This will require further study when additional populations
of M. vestita are located.

The berry measurements given in the species description are based on fruits that are not yet fully
mature. They are sure to increase somewhat when ripened fruits are collected.

ETYMOLOGY. — The specific epithet is derived from the Latin word vestitus, meaning clothed or
covered, alluding to the conspicuous hair covering on the abaxial foliar surface.

ACKNOWLEDGMENTS

I thank Robert L. Dressler for augmenting locality information for Miconia correae and
Mireya D. Correa for bringing several important collections to my attention. Logistical support was
provided by the Smithsonian Tropical Research Institute in Panama and the Museo Nacional de Costa
Rica and the Instituto Nacional de Biodiversidad in Costa Rica. I am also grateful to the herbaria cited
for gifts and/or loans of critical specimens and to the following individuals for preparing the line
drawings: Sheva Myers (Figure 1), Ellen del Valle (Figures 2-4), Jenny Speckels (Figures 5 and 7),
and Margaret de Jong (Figure 6).

54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 4

LITERATURE CITED

ALMEDA, F. 1981. The Mexican and Central American species of Adelobotrys (Melastomataceae). Ann. Mis-
souri Bot. Gard. 68:204—212.

. 1989a. Five new berry-fruited species of Tropical American Melastomataceae. Proc. Calif. Acad. Sci.

46(5):137—150.

. 1989b. New species and taxonomic notes on Mexican and Central American Melastomataceae. Proc.

Calif. Acad. Sci. 46(9):209—220.

. 1989c. Tessmannianthus, an arborescent genus of Melastomataceae new to Panama. Ann. Missouri Bot.

Gard. 76:16.

. 1990. New species and new combinations in Blakea and Topobea (Melastomataceae), with an historical
perspective on generic limits in the tribe Blakeeae. Proc. Calif. Acad. Sci. 46(14):299-326.

COGNIAUX, C. A. 1891. Mélastomacées. /n Monographiae phanerogamarum. Vol. 7. A. de Candolle and C. de
Candolle, eds. G. Masson, Paris. 1256 pp.

GLEASON, H. A. 1958. Melastomataceae. /n Flora of Panama, R. E. Woodson, Jr. and R. W. Schery, eds. Ann.
Missouri Bot. Gard. 45:203—304.

Lewis, W. H. 1971. High floristic endemism in low cloud forests of Panama. Biotropica 3:78—80.

STANDLEY, P. C. AND L. O. WILLIAMS 1950. New plants from Honduras. Ceiba 1:38-49.

WuRDACK, J. J. 1986. Atlas of hairs for neotropical Melastomataceae. Smithsonian Contr. Bot. 63:1—80

© CALIFORNIA ACADEMY OF SCIENCES, 2000
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San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

JUL 94 2000
Volume 52, No. 5, pp. 55-64, 4 figs., 3 tables, 2 plates. July 26, 2000

A New Species of Giant, Montane Phrynobatrachus (Anura:
Ranidae) from the Central Mountains of Kenya

by

Robert C. Drewes
Department of Herpetology, California Academy of Sciences
Golden Gate Park,San Francisco, California 94118

and

Jean-Luc Perret
Département d‘herpétologie et d ‘ichtyologie
Muséum d ‘Histoire naturelle, Route de Malagnou 1
CH-1211 Geneve 6, Switzerland

A new species of large forest-dwelling frog of the genus Phrynobatrachus Gunther
(Ranidae) is described, based on material collected in two different montane localities in
central Kenya. The new species differs in at least 10 characters from Phrynobatrachus
krefftii Boulenger, an endemic to the Eastern Arc Mountains of Tanzania, and the only
other large species of the genus in East Africa. A comparison of selected internal morpho-
logical character states among P. krefftii, the new species and several other species of the
genus Phrynobatrachus suggests that the new species and P. krefftii are not closely related.

The ranid genus Phrynobatrachus Ginther, 1862 is a poorly understood group. Individuals col-
lected in the field are often difficult to identify to species, and past workers have frequently confused
members of this group with cryptic, leaf-litter species of other genera including Arthroleptis,
Schoutedenella, Phrynodon and Dimorphognathus. The few external characters that distinguish
Phrynobatrachus from other genera include presence of a tarsal tubercle, absence of mandibular
odontoids (present in Dimorphognathus and Phrynodon) and absence of a median dorsal skin raphe
(present in Arthroleptis and Schoutedenella).

Of the 65 species of Phrynobatrachus listed in Frost (1985), many are based on scanty descrip-
tions. Because of their cryptic nature, few species exhibit obvious, easily recognizable field character-
istics upon which descriptions or identifications can be based; an exception is P. cricogaster (Perret
1957). Many species also exhibit a high degree of polymorphism (Stewart 1974). To date, few adver-
tisement calls of Phrynobatrachus species have been analyzed or published.

Herein, we describe a new species which is apparently endemic to the central mountain massif of
Kenya. This description is based on material collected at two separate localities between 20 and 30
years ago and deposited in four different institutions where it has remained unnoticed until recently.
Institutional abbreviations follow Leviton et al. (1985).

55

56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SPECIES DESCRIPTION

Phrynobatrachus irangi, n. sp.
Plate | and Fig. 1

HOLOTYPE. — MHNG 2230.76 male; KENYA: Meru District: Irangi Forest, above Irangi Forest
Station, springs of Ena Stream. 0°24’S, 37°28’E, elevation 2100 m. Collected by J.-L. Perret and
V. Mahnert, 12 October 1977.

PARATYPES. — MHNG 2230.74-75 males, MHNG 2230.77 sub-adult, MHNG 2230.78 female;
CAS 158967 male, CAS 158966 (male, cleared and stained); AMNH 68808, 68810-11, AMNH
72854-56 males; AMNH 68790, 68809, 72853 females: vicinity of type locality, collected by
R. Keith, 4 March, 28 April, 8 May, 1962 at 1965 m and 2286 m.

DIAGNOSIS. — A new montane species of Phrynobatrachus from the Kenya Massif and the
Aberdare Range differing from all East African congeners in the large size of the females (to
51.0 mm), and from the large Tanzanian Highlands species P. krefftii in the following external (1-6)
and internal (7-11) characters: (1) reduced webbing of the foot, (2) smaller inner metatarsal tubercle,
(3) longer fifth toe, (4) central plantar surface of foot in breeding males with spiny asperities, (5) snout
in breeding males rounded, not sharply projecting; (6) breeding males without dark outlining of lower
jaw and without chrome yellow gular coloration, (7) nasals widely separated, not greatly dilated medi-
ally, not overlapped posteriorly by sphenethmoids, (8) neural spines of vertebrae not strongly imbri-
cate, (9) base of omosternum not bifurcate, (10) base of thyrohyal originating well posterior to base of
posterolateral process of hyoid, and (11) sternal style semi-rectangular, not strongly tapered medially.

ETYMOLOGY. — The specific epithet, a noun in apposition, refers to the type locality, Irangi For-
est, Meru District, Kenya.

DESCRIPTION OF HOLOTYPE. — Male, 46.0 mm snout-vent length; habitus stout, robust; snout
protruding but rounded, not sharply angled (Fig. 2); diameter of eye 4.5 mm, slightly more than half
the distance to tip of snout, nearly equal to interorbital space; tympanum an oblique oval, its diameter
approximately three-fourths that of eye and situated beneath a dermal fold that originates from
mid-point of posterior margin of eye, curves ventrally around tympanum and terminates midway be-
tween angle of jaw and origin of forelimb; tip of fingers and toes rounded, only slightly dilated; tip of
each toe in males with one or two lateral spines; webbing between fingers absent, webbing between
toes reduced (Fig. 3) webbing formula 1[2-2+II1-2+III3-31V3+-2+V (Savage and Heyer 1997);
subarticular tubercles of both hands and feet single, palmar surfaces smooth, plantar surfaces beset
with numerous, small, pale-colored spinose asperities, especially conspicuous along axes of fourth
and fifth metatarsals (Fig. 3); large, thick greyish, granular nuptial pad extending from origin of
thumb to just beyond proximal subarticular tubercle; tarsal tubercle a pale-colored eminence capped
by white spine; fore- and hindlimbs stout, muscular; hindlimb length 2.5 times greater than snout-
vent length; inner metatarsal tubercle white, small, about one-third diameter of eye.

Dorsum generally smooth in appearance but beset with widely spaced, very small, white-pointed
tubercles that extend laterally to mid-lateral surface of body; thin, glandular ridge extends from the
posterior margin of each eye, angling medially to level above posterior extent of tympanum but not
converging with its fellow; a second, more posterior pair of slightly shorter, thin glandular ridges di-
verges obliquely in reverse direction to position above mid-point of forelimb insertion, forming fol-
lowing configuration:

al

i

dorsal skin of forelimbs smooth, hindlimbs smooth in femoral region but becoming increasingly tu-
berculate from midpoint of tibio-fibula to foot; posterior surface of thighs generally smooth with

DREWES AND PERRET: PHRYNOBATRACHUS IRANGI a7

PLATE |. A. (left) Holotype of Phrynobatrachus irangi (MHNG 2230.76). B. (right) Paratype (MHNG 2230.75). Photos by
JLP in type locality, Irangi Forest, Kenya.

white-tipped, small spinous tubercles interspersed with larger flattened warts; ventral surface of body
smooth, except for gular region, which consists of series of longitudinal, unpigmented, distensible
folds comprising the vocal pouch.

Color in preservative. Ground color a medium, muddy brown; pale areas darkish beige; ventrum
pale beige. Entire snout pale-colored, separated from darker dorsal coloration by sharply demarcated
line across interorbital space including anterior one-third of each eyelid; pale area extending
ventro-laterally and obliquely onto upper lip from anterior margin of eye; upper lip dark from poste-
rior margin of pale patch and below eye to anterior margin of tympanum; tympanum dark; rounded
pale patches extend antero-dorsally from anterior margin of origin of forelimbs to lateral margins of
anterior pair of glandular ridges, and distally onto anterior surfaces of forearms to point near wrists;
dorsal aspects of hands dark with darker band near each wrist; hindlimbs with nine, roughly
equally-spaced, dark bands extending from thighs to feet; ventrum immaculate.

Color in life (Plate 1A). Dorsum dark brown; entire snout light orange-brown, sharply demar-
cated by transverse line running between eyelids; large patch of same contrasting color on shoulder,
upper arm and elbow; hind limbs pale brown with dark, thick transverse bands; ventrum yellowish
tan; gular region somewhat grayish.

VARIATION IN PARATYPES. — Morphological features in the paratype series are generally con-
sistent with those described in the holotype. All female and a few male (AMNH 72854-72856) speci-
mens lack small, pale dorsal and lateral spines; these are present in all remaining male specimens.
Some variation exists in the plantar spines: plantar spines in the fourth metatarsal area are absent in the
largest females (AMNH 72853 and AMNH 68790), present but reduced in females AMNH 68809
and MHNG 2230.78; present but reduced in males AMNH 72854, 72855 and 72856. Males AMNH
72854-72856 are soft and rather poorly preserved in comparison with the rest of the type series in-
cluding males AMNH 68808, AMNH 68810-68811; these specimens are adults (snout-vent length
36.6—-45.7 mm) so absence of dorsal and lateral spines in males is probably an artifact of preservation.
Tarsal tubercles also vary; some individuals have a row of small tarsal spines, the largest and most
posterior of which also occupies the position typical of tarsal tubercles in other species in the genus.

All specimens in the type series are consistent in dorsal color pattern with minor variations in in-
tensity of ground color; all specimens exhibit the sharply-defined, pale-colored snout except one male

58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 5

paratype (MHNG 2230.75) in which
the snout is darker brown, but still de-
marcated by a pale orange-brown
transverse bar (Plate 1B). One male
specimen (AMNH 68808) has a thin,
mid-dorsal pale stripe extending from
the posterior margin of the pale snout
patch to the tip of the urostyle; one fe-
male (MHNG 2230.74) is darker and
more mottled in dorsal aspect than the
remaining specimens. Supratym-
panic, rounded pale patches are
discernable in six males (CAS
158967, MHNG 2230.74, AMNH
68808, 72854, 72855); these are less
obvious in females and the remaining
male specimens.

Ventral surfaces of three males
(MHNG 2230.78, AMNH 68780,
AMNH 68809) have diffuse greyish
mottling that extends posteriorly

Elevation range (ft): from the gular region to the underside

Se ies : ceed of the thighs; the underside in AMNH

7000 - 11000 72853 is immaculate; the venters of

|} above 11000 three males (AMNH 68811, 72855

Nairobi and MHNG 2230.77) are also mottled

but less so than in females; the ventral

surface of the hindlimbs in

male AMNH 72854 _ is moderately
mottled in pale brown.

FiGure 1. Type locality of Phrynobatrachus irangi sp. nov.: Irangi For- Measurements of the type series,
est, Mount Kenya, and location of Aberdare Range population (NMK and meristic comparisons with
specimens): former farm of Edna Oxtoby at Kimande, Kenya.

Phrynobatrachus krefftii are summa-
rized in Tables 1 and 2.

VOICE. — The advertisement call of Phrynobatrachus irangi has been recorded (Perret) from a
chorus of many males; evidently no single voice was close enough for sonographic analysis. The call
is rather loud and evidently emitted during the day only. It was described by R. Keith (in her field
notes, AMNH Archives) as “raugh-araugh-aaaaraugh-arararaugh-raraugh.”

COMPARISONS

External Morphology

The only other species of large Phrynobatrachus in East Africa, P. krefftii Boulenger, 1909, is en-
demic to the Eastern Arc Mountain system (Howell 1993). In Table 1, we have summarized meristic
data taken from the type series of P. irangi and 16 males and females of P. krefftii from Amani, East
Usambara Mountains, Tanzania (see Additional Material Examined). Whereas adult males of both
species are comparable in size, adult P. irangi females attain much larger snout-vent lengths; in fact,
female P. irangi may be the largest member of the genus Phrynobatrachus.

DREWES AND PERRET: PHRYNOBATRACAUS IRANGI oy)

Both male and female P. irangi have longer hindlimbs than
P. krefftii, but smaller inner metatarsal tubercles. The tarsal tu-
bercle of P. irangi is larger and more conspicuous than that of P.
krefftii, which tends to be a single, small, spinous eminence at
the terminus ofa curved skin fold that originates near the poste-
rior margin of the inner metatarsal tubercle. However, in some
specimens of both species a series of small tubercles are pres-
ent, with the largest occupying the posterior-most position. The
plantar surfaces of the fourth metatarsal are spinose in male P.
irangi, and smooth in P. krefftii. The presence of these small as-
perities on the ventral surfaces of the toes and feet, while differ-
ently distributed in these two species, appears to be a unique
character within the genus Phrynobatrachus. Specimens of P.
krefftii have more extensive webbing between the toes than do
P. irangi (Fig. 3; webbing formula for P. krefftii:
I1-1110-O1111-21'V2+-1V [Savage and Heyer 1997]).

Breeding male P. krefftii have a pointed, projecting, shale eee ish lateralleeneetsonPhrymo-
low snout in lateral profile (Fig. 2); the lower jaw is sharply de- batrachus irangi male holotype (MHNG
fined by dark pigment which, in turn, is outlined medially by 2230.76) (above); Phrynobatrachus kreff-
starkly contrasting white pigment. The gular region is bright /! Mle CAS (186541) (below).
chrome yellow in life (Barbour and Loveridge 1928). In preser-
vative, the yellow gular region of male P. krefftii fades to the same color as the venter, but the strongly
marked coloration of the lower jaw persists. Male P. irangi have rounded, less-projecting snouts, the
lower jaw is mottled, not evenly outlined, and the bright chrome yellow gular coloration of P. krefftii
males is absent.

TABLE |. Measurements of type series of Phrynobatrachus irangi. Mean (in mm) followed by range (in
parentheses).

n SVL Tibia T/SVL (%) Foot F/SVL (%) T/F (%)

P. irangi (male) 14 41.7 25.8 58.7 26.2 63.2 105.7
(36-46) (21-27) (54.7-62.3) = (21-29) = (53.4-64.0) = (93.1—108.6)
P. irangi (female) 4 45.7 24 Sh) 25.7 55.6 O5a1

(45-51) (22-26) (48.9-5S8.25) (24-28) (47.05-63.4) (88.5—108.3)

TABLE 2. Comparison of mean measurements (in mm) of Phrynobatrachus irangi and P. krefftii.

SVL Tibia T/SVL(%) Foot F/SVL(%) T/F(%) MT
P. irangi (male) 41.7 25.8 56.7 26.2 63.2 105.7 Weg)
P. krefftii (male) 41.0 21.8 53.4 24.2 59.4 89.9 2.4
P. irangi (female) 45.7 24.0 33.1 25.0 55.6 95.1 1.6

P. krefftii (female) 38.4 18.2 47.6 19.6 51.0 95:3 2.05

60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Internal Morphology

Examination of cleared and double-stained speci-
mens reveals notable differences between the two spe-
cies. In dorsal view, the nasals of P. irangi are widely
spaced, not greatly dilated medially and are not over-
lapped posteriorly by the sphenethmoids; P. krefftii
nasals are more broadly dilated medially, less widely
spaced, and overlapped by anterior projection of the
sphenethmoids (Plate 2A, B).

The presacral vertebrae in P. krefftii are strongly
imbricate; those of P. irangi are non-overlapping
(Plate 2A, B).

Viewed ventrally, the base of the omosternum in
P. irangi is slightly notched; that of P. krefftii is mod-
erately forked (see states 16.1 and 16.2, in Drewes
1984). The sternal style of P. irangi is near-
rectangular, only slightly compressed medially, with a
medial width greater than half the width of the anterior
margin; that of P. krefftii is compressed medially, so

ae eee that the medial width 1s half the width of the proximal
FIGURE 3. (left) Right foot in plantar view of male :
P. krefftii (NMK A/883.11); (right) P. irangi male Margin of the structure. (Plate 2C, D)
paratype (MHNG 2230.75). In P. irangi, the bases of the thyrohyals originate
posterior to a line drawn through the bases of the
posterolateral processes; in P. krefftii, the thyrohyals are deeply invasive into the corpus of the hyoid
plate and extend anterior to the bases of the posterolateral processes (Plate 2C, D)

In an attempt to ascertain the degree of relatedness between P. irangi and P. krefftii, the internal
characters that serve to separate the two were examined in cleared and stained specimens of eight ad-
ditional species of Phrynobatrachus (see Additional Specimens Examined). A phenogram, a
UPGMA tree generated by PAUP 4.0b1 (Fig. 4) was based on the character matrix in Table 3. An ad-
ditional state of character 1, degree of separation of nasals, was found in the added species, and coded
‘“M”. The phenogram, indicates that in spite of large size and gross similarity between P. irangi and P.
krefftii, they probably are not closely related. Phrynobatrachus irangi shares more of its diagnostic
character states with P. parvulus and P. parkeri than with the rest of the group; based on the same char-
acters, P. krefftii clusters with P. dendrobates.

The states of four characters (neural arch, omosternum, sternal style and separation of nasals)
were polarized using works by Clarke (1981), Drewes (1984), Lynch (1973) and a data set on
petropedetine ranid genera (Drewes, unpublished). PAUP was employed in an attempt to assess the
phylogenetic positions of P. krefftii and P. irangi with respect to each other and the relatively small
sample of additional species in the genus (15%, see Frost 1985). The resulting analysis yielded 311
most parsimonius trees and was not resolvable; however, it indicated that with respect to a presump-
tive ancestor, Phrynobatrachus irangi, P. kinangopensis, P. natalensis, P. parvulus, P. versicolor, P.
parkeri, P. perpalmatus, and P. plicatus form an unresolved polytomy, but are more closely related to
each other than any is to P. krefftii and P. dendrobates. The latter form a basal clade with respect to the
former. Interestingly, calling males of P. dendrobates share the characteristic bright, chrome yellow
gular region of P. krefftii males.

Both P. irangi and P. krefftii share spinous asperities in the subdigital areas of the feet, as well as
on the terminal discs, although the distribution of these asperities on plantar surfaces is consistently
different between the two species. To our knowledge, possession of this character state is unique to
these two species within the genus Phrynobatrachus; we posit that the appearance of this character

DREWES AND PERRET: PHRYNOBATRACHUS IRANGI

PLATE 2. Dorsal and ventral views of cleared and double-stained specimens of Phrynobatrachus krefftii (A, C, BM 1974.80)
and male paratype Phrynobatrachus irangi (B, D, CAS 158966). Key: 1. nasal, 2. sphenethmoid, 3. non-imbricate neural arch,
4. base of thyrohyal, 5. base of omosternum, 6. sternal style.

61

62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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TABLE 3. Distribution of internal character states differentiating P. irangi from P. krefftii among other
Phrynobatrachus species.

] a 3 4 5 6 fl
P. dendrobates M K K K K K K
P. kinangopensis I I I I K I K
P. natalensis (2) M K I I I ] I
P. parvulus I I I I I I I
P. versicolor ] K I I K K I
P. parkeri I ] I I I K I
P. perpalmatus I K K I K I I
P. plicatus (2) M K K I K K I
P. irangi ] I I I I I I
P. krefftii K K K K K K K

Character states: 1. nasal separation at midline

2. medial nasal dilation

3. posterior nasal overlap by sphenethmoid

4. neural arch imbrication (at least presacral vertebrae #1—3)

5. omosternum bifurcation

6. sternal style shape

7. thyrohyal condition

K = state found in P. krefftii

I= state found in P. irangi

M = state found in some additional species, i.e., nasals narrowly separated or meet at midline

may be the result of convergence, possibly related to their large size or derived from living in montane
habitats.

Natural history

Based on the field experience of Perret and R. Keith’s field notes, Phrynobatrachus irangi is ac-
tive by day. Males were found calling on banks of small streams in the forest, from under roots or logs,
and from holes in the mud. They were difficult to locate and capture. No amplexus has been observed,
and although tadpoles were seen in nearby clear water, their identity was not ascertained.

DISCUSSION

The history of the material upon which this description is based is interesting and also necessary
to document because of an error in the field data associated with the material from the National Mu-
seums of Kenya ([NMK], Additional Materials Examined). The specimens originally collected by E.
Oxtoby at Kimande were first brought to the attention of Drewes in the early 1970s by the then Head of
Herpetology at NMK. At that time, no attempt had been made on the part of NMK staff to collect fur-
ther material, in spite of urging on the part of Oxtoby. Following Oxtoby’s death, Drewes and an
NMK staffmember, Peter Nyamenya, made a day trip to the former Oxtoby property in October 1979
and found it severely modified by local subsistence farming. No Phrynobatrachus were seen or col-
lected. For the next several years, attempts on the part of Drewes to borrow the Oxtoby material from
the National Museums of Kenya for purposes of description were fruitless, and it was assumed that the
Oxtoby material had been misplaced or lost.

DREWES AND PERRET: PHRYNOBATRACHUS IRANGI 63

FiGuRE 4. UPGMA Phenogram generated by PAUP, based on character states in Table 3.

During a visit to the MHNG in Geneva in 1985, Perret showed Drewes a series of large
Phrynobatrachus that he and V. Mahnert had collected in the Irangi Forest in 1979. Irangi is on the
eastern slope of Mt. Kenya, while Kimande, the former Oxtoby property is on the south-southeastern
slope of the Aberdares, a range of mountains some 90 km., as the crow flies (see Fig. 1), from the
Irangi site. It became obvious to Drewes that if the two samples did not represent the same undes-
cribed species, they were almost certainly, closely related species. It seemed unwise to describe the
MHNG material without including the NMK material; attempts by both authors to borrow the NUK
material continued to be unsuccessful. In 1993, Drewes queried Dr. Linda Ford of the American Mu-
seum of Natural History as to whether or not Ronalda Keith, a field worker who made extensive col-
lections in East Africa in the early sixties, had ever collected in the Irangi Forest. The response was
positive and more than half of the type series described below is in the American Museum of Natural
History.

In 1993, during a visit to the NMK, Drewes learned that the original Oxtoby Aberdares material
had been found in a locked cabinet, and it has been included under Additional Material Examined.
With respect to the NMK material, it is clear that the cataloged collection date of “1979” is in error;
the specimens had to have been collected earlier, most likely in the early 1970s, inasmuch as the date
of the day trip undertaken by Drewes and Nyamenya in 1979 is documented in photographs of the
property taken at the time. The NMK specimens were not included as paratypes because the locality
from which they were taken has evidently been destroyed, the population may be extinct, and we
know of no field worker who has seen members of this population in life.

ADDITIONAL MATERIAL EXAMINED

Phrynobatrachus irangi: NMK A/1203/1-2, A/1203/4.Kenya: Murang’a Dist: Kimande. 0°49’S,
36°48'E; males, collected by E. Oxtoby, 30 April 1979(?); Kenya: Meru Dist: type locality: AMNH
68791-68793, AMNH 68795-68796, AMNH 135816. juveniles, collected by R. Keith, 1962.

Phrynobatrachus krefftii: CAS 168511-168557. Tanzania: Tanga Region: Muheza Dist: East
Usambara Mtns, vic. of Amani.; BMNH 1974.79-91. Tanzania: West Usambara Mtns. Mazumbai

64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 5

Forest Reserve, 4°48’S, 39°29’E; NMK A/883/7, A/883.11 males; A/883/16-17, females. Tanzania:
East Usambara: Amani collected by B. Watulege, July 1969.

Cleared and Stained: Phrynobatrachus dendrobates, CAS 145294: Zaire: Ituri Prov: Manguerete
Hipa; P. kinangopensis, CAS 152381: Kenya: Murang’a Dist: Kimande; P. krefftii, BM 1974.80:
Tanzania: West Usambara Mtns: Mazumbai Forest Reserve; P. natalensis, CAS 141564: Kenya:
Kakamega Dist: Kakamega Forest Station; CAS 141666: Kenya: Kakamega Dist: Lubao; P.
parvulus, CAS 145258: Zaire: Kivu Prov: Fizi Terr: Mokanga; P. versicolor, CAS-SU 13008: Zaire:
Kivu Prov: Albert N.P.: Kundhuru-ya-Tshuwe; P. parkeri, CAS 98168: Zaire: Uele: Monga; P.
perpalmatus, CAS 98156: Zaire: Uele: Albert N. P.: Buta; P. plicatus, CAS 136294, 136298: Ghana:
Eastern Reg.: Kade Agricultural Station.

ACKNOWLEDGMENTS

The authors are grateful to the following curators for loan of specimens and other material in their
care: Damaris Rotich and Alex Duff-MacKay (NMK), Barry T. Clarke (BM) and Linda Ford
(AMNH), and to Michelle Koo (CAS) for the map and Joseph Slowinski (CAS) for Figure 4.

LITERATURE CITED

BARBOUR, T. AND A. LOVERIDGE. 1928. A comparative study of the herpetological faunae of the Uluguru and
Usambara Mountains, Tanganyika Territory with descriptions of new species. Mem. Mus. Comp. Zool.,
Harvard Univ. 50:86—265, 4 pls.

CLARKE, B. T. 1981. Comparative osteology and evolutionary relationships in the African Raninae (Anura
Ranidae). Monit. zool. ital. (N. S.) Suppl. 15:285-331.

DREWES, R. C. 1984. A phylogenetic analysis of the Hyperoliidae (Anura): Treefrogs of Africa, Madagascar and
the Seychelles Islands. Occ. Pap. Calif. Acad. Sci. 139:1—70.

FROST, D. R., ed. 1985. Amphibian species of the world: a taxonomic and geographical reference. Assoc. Syst.
Collections and Allen Press, Lawrence, Kansas. v + 732 p.

HOWELL, K. M. 1993. Herpetofauna of the eastern African forests. Pp. 173-201 in Biogeography and ecology of
the rain forests of Eastern Africa. J. C. Lovett and S. K. Wasser, eds. Cambridge University Press, United
Kingdom.

LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology:
Part 1, Standard symbolic codes for institutional resource collections in Herpetology and Ichthyology.
Copeia 1985:802—832.

LYNCH, J. D. 1973. 3. The transition from archaic to advanced frogs. Pp.133-182 in Evolutionary biology of the
anurans. J. L. Vial, ed. University of Missouri Press, Columbia, Missouri.

PERRET, J.-L. 1957. Un nouveau Phrynobatrachus du Cameroun. Rev. suisse Zool. 46:527—5S31.

SAVAGE, J. M. AND W. R. HEYER. 1997. Digital webbing formulae for anurans: a refinement. Herpetol. Rev.
28:131.

STEWART, M. M. 1974. Parallel pattern polymorphism in the genus Phrynobatrachus (Amphibia, Ranidae)
Copeia 1974:823-832.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 6, pp. 65-75, 5 figs., 3 tables. JUL 34 2000 July 26, 2000

Two New Genera of Soft Corals (Anthozoa: Alcyoniidae) from
South Africa, with a Discussion of Diversity and Endemism in the
Southern African Octocorallian Fauna

by
Gary C. Williams

Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Two previously described taxa of soft corals, each apparently endemic to southern Africa,
are each given new generic names. The two species were originally described in the genus
Alcyonium Linnaeus, 1758, but are shown here to belong to distinct genera. Several south-
ern African soft coral taxa share a similar growth form, i.e., unbranched, digitiform to
capitate, with a conspicuously elongated stalk, and polyps restricted to a distal
polyparium. A comparison of superficially similar taxa is provided, along with a revised
faunistic analysis regarding diversity and endemism in the Octocorallia of southern
Africa.

Since 1985, a resurgence of interest in the systematics of southern African octocorals has pro-
duced a variety of publications describing various taxa comprising the fauna. Included here are:
Alderslade (1985), Benayahu (1993), Benayahu and Schleyer (1995, 1996), Branch, Griffiths,
Branch and Beckley (1994), Grasshoff (1988, 1991, 1992), Lopez-Gonzalez, Gili, and Williams
(2000), Ofwegen and Schleyer (1997), Richmond (1997), Verseveldt and Bayer (1988), Verseveldt
and Ofwegen (1992), Verseveldt and Williams (1988), Williams (1986a, 1986b, 1986c, 1987a,
1987b, 1988, 1989a, 1989b, 1989c, 1989d, 1990a, 1990b, 1992a, 1992b, 1992c, 1992d, 1992e, 1993,
in press), Williams and Lindo (1997), and Williams and Rogers (1989).

Two new genera of soft corals from the Cape Endemic Province of southern A frica (as defined by
Williams, 1992d), are here described. Each genus includes a single species previously described in the
genus Alcyonium. It is here argued that these taxa are better placed in genera other than Alcyonium.

METHODS

An examination of recently collected material was made for part of this study. The material was
collected by SCUBA and preserved in 70% ethanol. Sclerites were isolated using sodium
hypochlorite. Scanning electron micrographs were made on a Hitachi S-510 scanning electron micro-
scope. Photographic plates for publication were made using Adobe Photoshop 4.0 software. Abbrevi-
ations used in the text are as follows: CAS (California Academy of Sciences, San Francisco), CRRF
(Coral Reef Research Foundation, Palau), SAM (South African Museum, Cape Town).

65

66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 6

SYSTEMATIC ACCOUNT

Family Alcyoniidae Lamouroux, 1812

Lampophyton gen. nov.
non Alcyonium Linnaeus, 1758:803; Williams, 1992a:271 (in part).

DIAGNOSIS. — Upright, unbranched soft corals, clavate or torch-shaped. Stalk prominent, ex-
panding distally to form a flat-topped polyparium. Polyps monomorphic, retractile, restricted to the
distal terminus of the coral. Polyp calyces absent. Sclerites colorless, densely set coarse spindles,
more or less longitudinally arranged along the walls of the stalk and polyparium. Color alcohol
soluble.

TYPE SPECIES. — Alcyonium planiceps Williams, 1986.

ETYMOLOGY. — The new generic name is derived from the Greek, /ampas (a lamp or torch) and
phyton (a creature, either plant or animal); in reference to the torchlike appearance of colonies of this
species, with narrow stalk and expanded, flat-topped polyparium.

Lampophyton planiceps (Williams, 1986) new comb.
Figs. 1A—C, 2A-F, 3

Alcyonium planiceps Williams, 1986:53, figs. 1-7. 1992a:289, figs. 20 e-j, 21.

MATERIAL EXAMINED. CAS 118496 (Sta. No. SAFR 334), South Africa, Cape Province, off Port
Elizabeth, Algoa Bay, Riy Banks, 15—24 m, 23 February 1999; collected by John Starmer with aid of
SCUBA, two whole specimens. CAS 118497 (Sta. No. SAFR 365), South Africa, Cape Province, off
Port Elizabeth, Algoa Bay, White Sands 6, 20 m depth, 26 February 1999, collected by John Starmer
with aid of SCUBA, two whole specimens. Fig. 1B from SAM-H3280, Llandudno, Atlantic side of
Cape Peninsula, Cape Province, South Africa, 21 m in depth, 24 January 1984, collected by G. C.
Williams with aid of SCUBA. Fig. 1A, C from SAM-H3281, Llandudno, Atlantic side of Cape Penin-
sula, Cape Province, South Africa, 25-30 m in depth, 24 March 1984, collected by G. C.Williams
with aid of SCUBA.

DESCRIPTION. — The specimens examined range in length from 18—32 mm. In each specimen,
the distal polyparium arises from an upright and unbranched stalk, which in turn arises from the basal
holdfast. The distal-most portion of the stalk expands gradually to form the polyparium at the distal
terminus. This feature gives the body of the coral a club-shaped or torch-shaped appearance (Fig
1A—C). The polyps are restricted to the flat, disc-shaped distal terminus. The polyps are
monomorphic, retractile, and without calyces. The sclerites are relatively large and robust spindles
with varying amounts of surface ornamentation (Fig. 2A—F). The tubercles vary from simply rounded
and relatively sparsely distributed knoblike structures to more densely-set crownlike arrangements
(Fig. 2D-F). The sclerites are densely set and are mostly longitudinally arranged along the surface of
the stalk (Fig. 1A). A few sclerites are scattered in the shallow subsurface of the stalk and polyparium,
but are not present in the deep interior. The sclerites range in length from 0.82—1.80 mm. Sclerites are
altogether absent from the polyps. The wet preserved specimens have stalks that are tan to yellowish
brown in color (but apparently without zooxanthellae), with polyps and polyparies greenish gray or
brown. Pigments are alcohol soluble. The vivid coloration of living material is recorded in Fig. | A—C.

DISTRIBUTION. — Cape Province of South Africa; Atlantic side of the Cape Peninsula, Cape St.
Francis, Algoa Bay, and East London; 21—90 m in depth (Fig. 3).

WILLIAMS: TWO NEW GENERA OF SOFT CORALS 67

FiGuRE 1. Living soft corals A-C. Lampophyton planiceps. A. Colony with polyps retracted, total length 18 mm
(SAM-H3281). B. Colony with polyps partially expanded, total length 30 mm (SAM-H3280). C. Colony with polyps retracted,
total length 17 mm (SAM-H3281). D. Dimorphophyton mutabiliforme; 25 mm in length (SAM-H3716).

68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 6

REMARKS. — The surface of the stalk is often partly covered with a variety of epizoic organisms,
including tubicolous amphipods, tubicolous terebellid polychaetes, sponges, foraminiferans, and dia-
toms (Fig 1B, C).

Turbellarian flatworms and an undescribed species of the arminacean nudibranch genus
Dermatobranchus have also been found on colonies of Lampophyton planiceps. It is probable that the
nudibranch feeds on the soft coral. Both predator and prey exemplify similar rose lavender color-
ations.

Dimorphophyton gen. nov.

non Alcyonium Linnaeus, 1758:803; Williams, 1992a:271 (in part).

DIAGNOSIS. — Upright, unbranched soft corals, with changing growth form: digitiform or cylin-
drical in shape when expanded, capitate during retraction. Stalk conspicuous, comprising half of total
body length when expanded, more than half during retraction. Polyparium finger-shaped when ex-
panded, subspherical during retraction. Polyps monomorphic, retractile into the capitulum. Polyp ca-
lyces absent. Sclerites thin, flattened, irregularly-shaped rods with smooth surface texture,
tuberculation not evident. Sclerites colorless, few in number; sparsely scattered in anthocodiae and
often in circles surrounding the bases of the polyps. Rust orange color, alcohol-soluble.

TYPE SPECIES. — Alcyonium mutabiliforme Williams, 1988.

ETYMOLOGY. — The new generic name is derived from the Greek, di- (a prefix meaning two or
double), morphe (form or shape), and phyton (a creature, either plant or animal); in reference to the
two colony shapes (digitiform when expanded and capitate when retracted) displayed by these soft
corals.

Dimorphophyton mutabiliforme (Williams, 1988) new comb.
Figs? 1D; 2G IH: 3

Alcyonium mutabiliforme Williams, 1988:14, figs. 11-14; 1992a:286, figs. 19, 20 ad.

MATERIAL EXAMINED. — SAM-H3716, Hottentot’s Huisie, Atlantic side of Cape Peninsula,
Cape Province, South Africa, 15—18 m in depth, | 1 November 1984, four whole specimens, collected
by G. C. Williams with aid of SCUBA.

DESCRIPTION. — Williams (1988:14—19 and 1992a:286—289) has previously described this spe-
cies. A brief redescription is provided as follows: The seven known specimens are upright and un-
branched, each less than 40 mm in length. The distal polyparium arises from a conspicuous stalk,
which in turn arises from the basal holdfast. Expanded specimens are digitiform, with the cylindrical
polyparium comprising 50-55% of the total length of the specimen (Fig. 1D). Retracted specimens
are capitate, with the subspherical polypary comprising 35-45% of the total length of the specimen.
The monomorphic polyps are retractile and are evenly distributed over the surface of the polyparium.
They do not have calyces. Sclerites are very thin, relatively smooth, irregularly-shaped rods, with lit-
tle to no surface ornamentation. They vary in length from 0.06—0.26 mm (Fig. 2G, H). The sclerites
are very few innumber and are sparingly distributed in the anthocodiae; as well as on the surface of the
polyparium where they are arranged in rings surrounding the bases of most of the polyps. Sclerites are
absent from other areas of the specimens. The specimens are deep orange to rust orange in life (appar-
ently without zooxanthellae), fading to a dull reddish brown wet preserved. Pigments are alco-
hol-soluble. The vivid coloration of living material is recorded in Fig. 1D.

DISTRIBUTION. — Llandudno and Hottentot’s Huisie, Atlantic side of the Cape Peninsula, Cape
Province of South Africa; 15—21 m in depth (Fig. 3).

WILLIAMS: TWO NEW GENERA OF SOFT CORALS 69

FIGURE 2. Scanning electron micrographs of sclerites. A-F. Lampophyton planiceps. A. 1.18 mm in length. B. 0.82 mm in
length. C. Detail of one end ofa sclerite; length of portion shown 0.64 mm. D. Detail ofa single tubercle from sclerite shown in
C; scale bar = 0.005 mm. E. 1.20 mm in length. F. 1.20 mm in length. G-H. Dimorphophyton mutabiliforme. G. 0.16 mm in
length. H. 0.18 mm in length.

70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 6

Durban. indian
Ocean

31

East London,”

$ Cape Town Port Elizabeth. Maca bay
is

i”
Cape * Cape St. Francis

Cape Agulhas
21 25

Peninsula

FIGuRE 3. Map of southern Africa showing collecting stations for Dimorphophyton mutabiliforme (@) and Lampophyton
planiceps (@).

REMARKS. — The surface of the stalk is often partly covered with a variety of epizoic organisms
such as sponges, barnacles, and tunicates (Fig 1D).

DISCUSSION

SYSTEMATICS. — I have discussed and revised the definition of the genus A/cyonium Linnaeus,
1758, in earlier publications (1986a:262—264; 1986b:61—62; 1988:18—19; 1992a:271), and I also pro-
vided (1986:259-262) a synopsis of the history of taxonomy relating to this genus and related taxa.
Since that period, it has been generally agreed by workers in soft coral systematics (P. Alderslade and
L. P. van Ofwegen, pers. comm., and the present work) that it is more realistic to recognize a sensu
stricto definition reflecting the lobate or digitate growth form of the type secies A/cyonium digitatum
Linnaeus, 1758, rather than a broader sensu /ato definition encompassing a variety of growth forms as
I have listed previously (1988:19). It is proposed that as our knowledge of alcyoniid taxonomy in-
creases, species previously included in the genus A/cyonium that do not suitably fit the original defini-
tion, should either be placed in other previously described genera such as Eleutherobia Piitter, 1900,
or new genera should be named to accommodate them (such as Lampophyton gen. nov., and
Dimorphophyton gen. nov.).

In addition, recent study has indicated that the genus 4/cyonium as presently recognized may ac-
tually be comprised of two distinct clades: one representing cold-water to temperate, azooxanthellate,
often brightly-colored species such as Alcyonium fauri J. S. Thomson, 1910; and the other repre-
sented mostly by coral reef taxa of tropical regions, such as Alcyonium utinomii Verseveldt, 1971,
which are zooxanthellate and mostly white or light brown in color (pers. observ. and P. Alderslade,

WILLIAMS: TWO NEW GENERA OF SOFT CORALS 71

pers. comm.). It remains to be determined whether or not the latter clade deserves separate generic
designation. Alderslade (in press) addresses this issue.

Other taxa of southern African or Indian Ocean soft corals are superficially similar in growth
form (unbranched: digitiform, clavate, or capitate) to Lampophyton planiceps and Dimorphophyton
mutabiliforme, but differ as follows. Malacanthus capensis (Hickson, 1900), Acrophytum claviger
Hickson, 1900, Anthomastus spp., Paraminabea spp., and Verseveldtia spp. are dimorphic.
Bellonella spp. have spindles less than 0.15 mm in length, while Eleutherobia spp. have radiates.
Alcyonium spp. are mostly lobate to digitate or membranous in growth form. Pieterfaurea spp. have a
palisade-like arrangement of sclerites around the base of each polyp.

DIVERSITY AND ENDEMISM IN THE SOUTHERN AFRICAN OCTOCORALLIA. — I have previously
(1992d) provided a detailed biogeographic analysis of the octocorallian fauna of southern A frica, de-
scribing diversity and levels of endemism in the Cape Endemic Province, as well as faunal contribu-
tions to the regional biota from neighboring zoogeographic provinces. Included here are the
Indo-Pacific, Atlantic, and Southern Oceans (Antarctic and Subantarctic). Since 1992, new informa-
tion has allowed for a refinement of analysis concerning diversity and endemism in the fauna. Figures
4 and 5 summarize the results of this revised assessment. The charts are based on data presented in Ta-
bles 1-3. A revised list of valid species of southern A frican octocorals will be published after the pub-
lication of several generic revisions and descriptions of new taxa.

Some definitions relevant to the reassess-
ment are as follows: Soft corals are octocorals
without an internal axis (families Alcyoniidae,
Nephtheidae, Nidaliidae, and Xeniidae). Genus Number of species
Gorgonians refer to octocorals with an internal

TABLE |. Southern African Octocorallia - largest
genera by number of species.

axis (families Anthothelidae, Melithaeidae, pmlaria .
Keroeididae, Acanthogorgiidae, Plexauridae, 4) ..,onium 6
Gorgoniidae,_ Ellisellidae, Chrysogorgiidae, Sarcophyton a
Primnoidae, and Isididae). Widespread refers to Leptogorgia 5
scattered or cosmopolitan distributions as | have — Lobophytum 5
previously defined (1992d:356—357). This refer- Pieterfaurea 5

sli Simpsonella 5
ence also defines the limits of other eres 5
biogeographic regions such as Cape Endemic, Drifa A
Indo-Pacific, Atlantic, and Southern Oceans. 1 Virgularia 4
have previously elucidated the geographical lim- = Cavernularia 3
its of southern Africa (1992d:355). A particular Sale y

j 5 1 1 | 1 itypic genera

genus is assigned a biogeographic affinity if the Wena oe as

majority of species within the genus are endemic
to that particular biogeographic region (Wil- Tota] 165
liams 1992d:368—369).

TABLE 2. Southern African Octocorallia - biogeographic composition.

Biogeographic category Number of genera Number of species
Indo-Pacific 36 47
Widespread 30 9
Endemic 13 78
Atlantic 9 2
Antarctic Z 0

29

Unidentified -

Total 90 165

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 6

ANTARCTIC
inularia

Eleutherobia

Icyonium
monotypic genera

/ Sarcophyton

Leptogorgia

INDO-PACIFIC

obophytum

Pieterfaurea
J
impsonella

q

enia

ditypic genera

WIDESPREAD

>

Virgularia
Cavernularia
Eunicella

WIDESPREAD

UNDESCRIBED

ATLANTIC

WIDESPREAD

FiGure 4. Faunistic analysis of southern A frican Octocorallia. A. Largest genera by number of species per genus, n = 165 (es-

timated number of species). B. Biogeographic affinities of the genera, n = 90 (number of genera). C. Biogeographic categories

of species considered valid, n= 135 (number of valid described species). Biogeographic categories including undescribed spe-
cies, n = 165 (estimated number of species).

Of the thirteen largest genera in the fauna, nine are soft corals, two are gorgonians (Leptogorgia
and Simpsonella), and two are pennatulaceans (Cavernularia and Virgularia). The fauna also con-
tains 15 ditypic genera and 62 monotypic genera. The largest genus is Sinularia (9.1% of the fauna),
followed by Eleutherobia (4.3%), and Alcyonium and Sarcophyton (both 3.6%) (Table | and Fig. 4A).

An analysis of biogeographic affinities at the generic level shows that the Indo-Pacific represents
the largest source contribution to the fauna with 40%, followed by those with widespread distributions
(33%), and southern African endemic genera (15%). Atlantic (10%) and Antarctic or Southern Ocean
(2%) contributions are relatively minor (Table 2 and Fig. 4B). The previous assessment (Williams
1992d:369, fig. 12A) showed similar percentages, except for the Indo-Pacific (45%) and the endemic

component (10%). This change partly reflects the fact that four new endemic genera have been recog-
nized since the last analysis was published.

Regarding identified species that are considered valid, the endemic component is by far the larg-
est with 57%, followed by the Indo-Pacific with 35%, species with widespread distributions (7%), and

WILLIAMS: TWO NEW GENERA OF SOFT CORALS aS

Atlantic species (1%) (Table 2 and Figure 4C).
When the undescribed species that are consid-
ered valid (18%) are added to this, the endemic
component shows a contribution of 46%, while
the Indo-Pacific component represents 29%,
with minor contributions from the widespread
category (6%) and the Atlantic (1%) (Table 3
and Figure 4D).

It is presumed that if undescribed species
were described and added to the data set, that the
percentages shown in Figure 4C would probably
reflect a relatively accurate and consistent quan-

80
7
: _
rr)
®
rE
®
[ok
”
re)
~ 40
®
fe]
=
=|
za

soft corals

gorgonians

TABLE 3. Southern African Octocorallia -
comparative species richness.

Taxonomic group Number of species

Heliporaceans l

sea pens

Stoloniferans 9

Soft corals 78
Gorgonians 43
Pennatulaceans 34
Total 165

helioporaceans

stoloniferans

corals
ty oe

Mo PPP eee ee ff
stoloniferans helioporaceans

Octocorallian groups

FiGure 5. Species richness in southern African octocorals (n=1 65).

74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 6

titative portrayal of the faunistic elements. Williams (1992d:632, fig. 6F) shows a 53.30% contribu-
tion for endemics based on estimated number of species of Octocorallia. The new assessment shows a
56.72% contribution. Therefore, a reasonable interpretation is that over one-half of the southern A fri-
can octocorallian fauna is represented by endemic species.

Soft corals comprise the largest group regarding species richness (47%), followed by gorgonians
(26%), pennatulaceans (21 %), stoloniferans (5%), and helioporaceans (1%) (Table 3 and Figure 5).

ACKNOWLEDGMENTS

I am grateful to Phil Alderslade (Museum and Art Gallery of the Northern Territory, Darwin),
Yehuda Benayahu (Tel Aviv University, Tel Aviv), and Katie Martin (Scientific Publications, Cali-
fornia Academy of Sciences, San Francisco) for their comments and suggestions; and to John Starmer
(Coral Reef Research Foundation, Palau) for recently collected material.

LITERATURE CITED

ALDERSLADE, P. 1985. Redescription of Acrophytum claviger (Coelenterata: Octocorallia). The Beagle
2(1):105—113.

. Inpress. Three new genera of soft corals (Coelenterata: Octocorallia) with notes on the classification of
some established taxa. Zoologische Mededelingen.

BENAYAHU, Y. 1993. Corals of the South-west Indian Ocean I. Aleyonacea from Sodwana Bay, South Africa.
South African Association for Marine Biological Research, Oceanographic Research Institute, Investiga-
tional Report No. 67:1—16.

BENAYAHU, Y. AND M. H. SCHLEYER. 1995. Corals of the South-west Indian Ocean II. E/eutherobia aurea spec.
nov. (Cnidaria, Alcyonacea) from deep reefs on the KwaZulu-Natal Coast, South Africa. South African As-
sociation for Marine Biological Research, Oceanographic Research Institute, Investigational Report No.
68:1-12.

. 1996. Corals of the South-west Indian Ocean III. Alcyonacea (Octocorallia) of Bazaruto Island, Mo-
zambique, with a redescription of Cladiella australis (Macfadyen, 1936) and a description of Cladiella
kashmani spec. nov. South African Association for Marine Biological Research, Oceanographic Research
Institute, Investigational Report No. 69:1—22.

BRANCH, G., C. L. GRIFFITHS, M.L. BRANCH, AND L. E. BECKLEY. 1994. Two oceans, a guide to the marine life of
southern Africa. David Philip, Cape Town. 360 pp.

GRASSHOFF, M. 1988. The genus Leptogorgia (Octocorallia: Gorgoniidae) in West Africa. Atlantide Reports
14:91-147.

. 1991. Die von EE. J. C. Esper 1788-1809 beschriebenen Anthozoa (Cnidaria). 1. Die Sammlung Esper im

Senckenberg-Museum. II. Octocorallia. II]. Antipatharia. Senckenbergiana biol. 71(4/6):325—368.

. 1992. Die Flachwasser-Gorgonarien von Europa und Westafrika (Cnidaria, Anthozoa). Courier
Forschungsinstitut Senckenberg 149:1—135.

LOPEZ-GONZALEZ, P., J.-M. GILI, AND G. C. WILLIAMS. 2000. On some veretillid pennatulaceans from the east-
ern Atlantic and western Pacific Oceans (Anthozoa: Octocorallia), with a review of the genus Cavernularia
Valenciennes, and descriptions of new taxa. Journal of Zoology 250(2):201—216.

OFWEGEN, L. P. VAN AND M. H. SCHLEYER. 1997. Corals of the South-west Indian Ocean V. Leptophyton
benayahui gen. nov. and spec. nov. (Cnidaria, Alcyonacea) from deep reefs off Durban and on the
KwaZulu-Natal south coast, South Africa. South African Association for Marine Biological Research,
Oceanographic Research Institute, Investigational Report No. 71:1—12.

RICHMOND, M. D., ed. 1997. A guide to the seashores of eastern African and the western Indian Ocean islands.
Sida, Department for Research Cooperation, SAREC, Stockholm, Sweden. 448 pp.

VERSEVELDT, J. AND F.M. BAYER. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and
Nidaliopsis (Octocorallia: Alcyoniidae and Nidaliidae), with descriptions of two new genera. Zoologische
Verhandelingen 245:1—131.

VERSEVELDT, J. AND L. P. VAN OFWEGEN. 1992. New and redescribed species of A/cvonium Linnaeus, 1758
(Anthozoa: Alcyonacea). Zoologische Mededelingen Leiden 66 (7):155—181.

WILLIAMS: TWO NEW GENERA OF SOFT CORALS 1

VERSEVELDT, J. AND G. C. WILLIAMS. 1988. A redescription of the soft coral Alcyonium valdiviae Kukenthal,
1906, with the description of a new species of Litophyton Forsskal, 1775 from southern Africa
(Octocorallia, Alcyonacea). Annals of the South African Museum 97(12):3 15-328.

WILLIAMS, G. C. 1986a. Morphology, systematics, and variability of the southern African soft coral Alcyonium
variabile (J. Stuart Thomson, 1921) (Octocorallia, Alcyoniidae). Annals of the South African Museum
96(6):241-270.

. 1986b. A new species of the octocorallian genus A/cyonium (Anthozoa: Alcyonacea) from southern Af-

rica, with a revised definition of the genus. Journal of Natural History 20(1):53—63.

. 1986c. What are corals? Sagittarius, Natural History Magazine of the South African Museum, Cape

Town 1(2):11-15. [Reprinted in Underwater, Ihlane Publications (Pty) Ltd., Northway, Natal, South Africa

6:26—27].

. 1987a. A new species of stoloniferous octocoral (Cnidaria: Alcyonacea) from the southwestern Indian

Ocean. Journal of Natural History 21(1):207—218.

. 1987b. The aberrant and monotypic soft coral genus Malacacanthus Thomson, 1910 (Octocorallia:

Alcyontidae), endemic to southern Africa. Journal of Natural History 21(6):1337—1346.

. 1988. Four new species of southern African octocorals (Cnidaria: Aleyonacea) with a further diagnostic

revision of the genus A/cyonium Linnaeus, 1758. Zoological Journal of the Linnean Society 92(1):1—26.

. 1989a. A review of recent research on the sublittoral coral reefs of northern Natal with a provisional as-

sessment of findings regarding the distribution of octocorals on Two-Mile Reef, Sodwana Bay. South A fri-

can Journal of Science 85(3):140—-141.

. 1989b. A provisional annotated list of octocorallian coelenterates occurring on the sublittoral coral reefs

at Sodwana Bay and Kosi Bay, northern Natal, with a key to the genera. South African Journal of Science

85(3):141-144.

. 1989c. The pennatulacean genus Cavernularia Valenciennes (Octocorallia: Veretillidae). Zoological

Journal of the Linnean Society 95(4):285—310.

. 1989d. A comparison of the stoloniferous octocorallian genera Bathytelesto, Stereotelesto, Rhodelinda,

and Scyphopodium, with the description of a new species from southeastern Africa (Anthozoa:

Clavulariidae). Journal of Zoology 219(4):621—635.

. 1990a. A new genus of dimorphic soft coral from the southwestern fringe of the Indo-Pacific

(Octocorallia: Alcyoniidae). Journal of Zoology 221(1):21-35.

. 1990b. The Pennatulacea of southern A frica (Coelenterata, Anthozoa). Annals of the South African Mu-

seum 99(4):3 1-119.

. 1992a. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals (Coelenterata,

Anthozoa). Annals of the South African Museum 100(3):249-358.

. 1992b. The Alcyonacea of southern Africa. Gorgonian octocorals (Coelenterata, Anthozoa). Annals of

the South African Museum 101(8):181—296.

. 1992c. Revision of the Indo-Pacific soft coral genus Minabea Utinomi, 1957, with new taxa from the

Indo-West Pacific. Proceedings of the California Academy of Sciences 48(1):1—26.

. 1992d. Biogeography of the octocorallian coelenterate fauna of southern Africa. Biological Journal of

the Linnean Society 46(4):35 1-401.

. 1992e. Revision of the gorgonian genus Simpsonella (Octocorallia: Chrysogorgiidae) from the western

margin of the indo-Pacific, with the description of a new species from southeastern A frica. Zoological Jour-

nal of the Linnean Society 105:377-405.

. 1993. Coral reef octocorals: an illustrated guide to the soft corals, sea fans, and sea pens inhabiting the

coral reefs of northern Natal. Durban Natural Science Museum, Durban, South Africa. 64 pp.

.In press. A review of the endemic southern African soft coral genus Pieterfaurea (Octocorallia:
Nidaliidae), with descriptions of three new species. Zoologische Mededelingen, Leiden.

WILLIAMS, G.C. AND K. LINDO. 1997. A review of the octocorallian genus Leptogorgia (Anthozoa:
Gorgoniidae) in the Indian Ocean and subantarctic, with description of a new species and comparisons with
related taxa. Proceedings of the California Academy of Sciences 49(15):499-521.

WILLIAMS, G. C. AND J. ROGERS. 1989. Photographic evidence of bathyal octocorals from the Cape Basin. South
African Journal of Science 85(3):191—192.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 7, pp. 77-85, 5 figs. July 26, 2000

Redescription and Reassessment of Cadlina luarna
(Ev. Marcus and Er. Marcus, 1967), comb. nov. (Mollusca,
Opisthobranchia, Doridina)

by

Angel Valdés
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Orso Angulo Campillo
Departamento de Biologia Marina, Laboratorio de Plancton,
Universidad Autonoma de Baja California Sur, A.P. 19-B,
La Paz, Baja California Sur, México 23080

Inuda luarna is the type species of the problematic genus /nuda, and the only species so
far assigned to it. Several newly collected specimens from Baja California Sur, Mexico, al-
lowed a redescription of this species. These specimens are clearly conspecific with the origi-
nal type material of Jnuda luarna, which was also re-examined, but show some anatomical
differences. The external morphology of the living animals of this species, including the
presence of mantie glands, is described for the first time.

The anatomical features of Jnuda luarna are very similar to those of the genus Cadlina,
and /nuda is regarded as a junior synonym of Cadlina. Cadlina luarna appears to be a basal
member of the Cadlina clade, and retains several plesiomorphies also present in
Actinocyclus.

RESUMEN

Inuda luarna es \a especie tipo del problematico género /nuda, y la unica especie que le ha
sido asignada hasta este momento. Varios ejemplares recolectados en Baja California Sur,
México, han permitido redescribir esta especie. Estos especimenes son claramente
conespecificos con el material tipo de Jnuda luarna, que también ha sido re-examinado,
pero presentan algunas diferencias anatomicas. La morfologia externa de los animales
vivos, incluyendo la presencia de glandulas del manto, se describe por primera vez.

Las caracteristicas anatémicas de Inuda luarna son muy similares a las del género
Cadlina, e Inuda es considerado como un sin6énimo de este ultimo. Aparentemente Cadlina
luarna es un miembro basal de Cadlina, y mantiene varias caracteristicas plesiomorficas
que también estan presentes en Actinocyclus.

The genus /nuda was originally described by Marcus and Marcus (1967) based on a single spe-
cies, Inuda luarna, which is the type species by original designation. Since then, no additional species
have been assigned to the genus /nuda, and the systematic position of this taxon remained uncertain.

The anatomy of /nuda was studied by Marcus and Marcus (1967), but the information provided
was not adequate by modern standards. In addition, the external morphology and coloration of the liv-
ing animals were unknown.

iG)

78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No.7

Most authors agreed to consider /nuda as a member of the Chromodorididae (Marcus and Marcus
1967; Skoglund 1991; Angulo Campillo 2000), except for Keen (1971), who placed this genus in its
own family-level taxon. However, Rudman (1984), in his review of the genera of Chromodorididae,
did not refer to /nuda.

This paper redescribes /nuda luarna based on newly collected material from Baja California Sur,
Mexico, and attempts to determine the relationships of this taxon.

The material examined is deposited in the Department of Invertebrate Zoology and Geology of
the California Academy of Sciences (CASIZ), the Museo de Historia Natural de la Universidad
Autonoma de Baja California Sur (MHNUABCS) and the National Museum of Natural History,
Washington D. C. (USNM).

SPECIES DESCRIPTION
Genus Cadlina Bergh, 1879

Acanthochila Mérch, 1869, suppressed by Opinion 812 (ICZN, 1967). Type species: Doris laevis Linnaeus,
1767.

Echinochila Morch, 1869, suppressed by Opinion 812 (ICZN, 1967). Type species: Doris laevis Linnaeus, 1767.

Cadlina Bergh, 1878 (nomen nudum).

Cadlina Bergh, 1879. Type species: Doris repanda Alder and Hancock, 1842 (= Cadlina laevis Linnaeus, 1767),
by original designation.

Juanella Odhner, 1922. Type species: Juanella sparsa Odhner, 1922, by monotypy.

Inuda Ev. Marcus and Er. Marcus, 1967. Type species: /nuda luarna Ey. Marcus and Er. Marcus, 1967, by origi-
nal designation (new synonym).

Cadlina luarna (Ev. Marcus and Er. Marcus, 1967)
Figs. 1-5

Inuda luarna Marcus and Marcus, 1967:182—184, figs 38-44; Keen, 1971:826; Abbott, 1974:356; Farmer,
1980:104; Skoglund, 1991:12; Gonzalez, 1993:247; Angulo Campillo, 93—94, fig. 41.

TYPE MATERIAL. — LECTOTYPE (here selected): Puerto Penasco, Sonora, Mexico, 23 mm pre-
served length, contracted and dissected, leg. P. Pickens (USNM 678405).

REMARKS ON THE TYPE MATERIAL. — There is a single specimen of /nuda luarna deposited at
USNM.. It was collected from Puerto Penasco, Sonora, Mexico (type locality) by P. Pickens, and has
several labels. One of the labels, handwritten by Eveline Marcus, only includes the name of the spe-
cies. Another label, handwritten by a different person (probably the collector) contains the numbers
“14.22.” A more modern label, printed with USNM format indicates the name of the species, locality,
collector and that this specimen is the holotype of /nuda luarna. The two remaining labels (also
USNM format) just repeat the taxonomic status and the registration number of the specimen.

Marcus and Marcus (1967) mentioned that they examined two specimens for the description of
this species, and did not select either of them to be the holotype. Thus, the specimen deposited at the
USNM collection is not the holotype, but one of the two syntypes of /nuda luarna. Since the other
specimen is untraceable, and its identity could not be confirmed, we designate the available syntype as
the lectotype of this species.

ADDITIONAL MATERIAL EXAMINED. — Calerita, Bahia de La Paz, Baja California Sur, Mexico,
23 February 1997, 2 specimens 55—60 mm long, leg. O. Angulo Campillo(MHNUABCS-INV 1808).
Calerita, Bahia de La Paz, Baja California Sur, Mexico, 10 April 1999, 3 specimens 20-44 mm pre-
served length, leg. O. Angulo Campillo (CASIZ 121103). Ensenada de los Muertos, southeast of La

VALDES AND ANGULLO CAMPILLO: REDESCRIPTION OF CADLINA LUARNA

54

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FiGure 1. Living animal of Cadlina luarna from Baja California Sur, Mexico (CASIZ 121104). A. Dorsal view; B. Detail of

the gill; C. Detail of the border of the mantle showing mantle glands. Abbreviations: a, anal papillae; mg, mantle gland.

80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No.7

FIGURE 2. Scanning electron micrographs of Cadlina luarna (CASIZ 121104). A. Dorsal surface, scale bar = 600 ppm. B. De-
tail of a mantle gland, scale bar = 100 pm. Abbreviations: mg, mantle gland; tb, tubercle.

Paz, Baja California Sur, Mexico, 11 March 2000, 1 specimen 42 mm preserved length, leg. O.
Angulo Campillo (CASIZ 121104). Mazatlan, Sinaloa, Mexico, 8 December 1932, 1 specimen
36 mm preserved length, collector unknown (CASIZ 073360).

EXTERNAL MORPHOLOGY. — The body shape is oval to rounded (Fig. 1 A). The center of the dor-
sum is elevated. There are numerous, low and rounded tubercles (Fig. 2A) that are more densely con-
centrated on the mantle margin. On the mantle margin there are several rows of small mantle glands
(Figs 1C, 2B). They are conical in shape and irregularly arranged. The gill is composed of nine bipin-
nate branchial leaves (Fig. 1B). The perfoliate rhinophores have 12 lamellae. The anal papillae is situ-
ated in the center of the circlet of branchial leaves (Fig. 1B).

The foot sole is narrow, about half as wide as the mantle margin. The anterior border of the foot is
grooved but not notched. The oral tentacles are short and wide, having a deep notch on their ventral
side (Fig. 3G).

The background color of the body is pale brown. There are numerous pale creamy white, rounded
blotches, that are more densely arranged on the mantle margin. These blotches may be fused together
forming larger creamy white areas. The center of the dorsum is covered with minute dark brown spots.
The branchial sheath is pale creamy white, and the rhinophoral sheaths translucent gray. The gill and
rhinophores are dark brown, with the rachis translucent pale gray. The mantle glands are bright
orange.

ANATOMY. — Digestive system. The oral tube has three strong muscles that attach to the body
wall (Fig. 3D). The muscular buccal bulb is about four times shorter than the oral tube and has two ad-
ditional muscles attached. The jaws are composed of a number of bifid elements, about 20 um long
(Fig. 4D). The radular formula is 135 x 65.1.65 in one specimen examined (CASIZ 121104). The
rachidian teeth have a single central cusp, and one or two large denticles on each side (Fig. 4A). The
mid-lateral teeth have a strong, short cusp, and one or two triangular denticles on the outer side (Fig.
4B). The outermost teeth are elongate, having one to three small denticles (Fig. 4C). The esophagus

VALDES AND ANGULLO CAMPILLO: REDESCRIPTION OF CADLINA LUARNA 81

FiGuRE 3. Anatomy of Cadlina luarna. A. Dorsal view of the internal organs (CASIZ 121104), scale bar = 2 mm; B. Repro-
ductive system (CASIZ 121104), scale bar = 1 mm; C. Detail of several reproductive organs (CASIZ 121104), scale
bar = 1 mm; D. Central nervous system (CASIZ 121104), scale bar = 1 mm; E. Lateral view of the anterior portion of the diges-
tive system (CASIZ 121104), scale bar = 1 mm; F. Penial hooks (CASIZ 121104), scale bar = 10 um. G. Ventral view of the
mouth area (CASIZ 121103), scale bar = 5 mm. Abbreviations: ag, abdominal ganglion; am, ampulla; at, genital atrium; b,
blood gland; bb, buccal bulb; bc, bursa copulatrix; bg, buccal ganglion; c, cerebral nerve; cg, cerebral ganglion; dd, deferent
duct; dg, digestive gland; eg, esophageal ganglion; es, esophagus; fg, female glands; h, heart; i, intestine; m, retractor muscle; 0,
oral tentacle; ot, oral tube; p, pedal nerve; pc, pedal commissure; ppc, parapedal commissure; pg, pedal ganglion; pl, pleural
nerve; plg, pleural ganglion; pr, prostate; r, rhinophoral nerve; rg, rhinophoral ganglion; rs, radular sac; sg, salivary gland; sh,
syrinx; sr, seminal receptacle; st, stomach; v, vagina; vg, vestibular gland; vl, visceral loop.

82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No.7

opens into the proximal end of the buccal bulb. Near this point two large salivary glands attach to the
buccal bulb. The esophagus is short, connecting distally to the digestive gland. The stomach 1s oval
and connects distally to the long intestine, which runs almost straight down to the anal opening
(Fig. 3A).

Reproductive system. The ampulla is very long and convoluted (Fig. 3B). It branches into the
short oviduct and the prostate. The prostate is short and tubular, but well differentiated. It narrows into
the deferent duct, which expands again into the muscular ejaculatory portion. The penis is armed with
several rows of hooks (Fig. 3F). These hooks have an elongate cusp about 20 um long and a shorter
base. There is an undifferentiated vestibular gland near the distal aperture of the female glands. The
vagina is long and convoluted. At mid-length it branches into a duct that connects to the seminal re-
ceptacle and the uterine duct (Fig. 3C). The uterine duct is short and thin, and opens near the center of
the female gland mass. The seminal receptacle is slightly oval, almost as large as the pyriform bursa
copulatrix.

Central nervous system. The cerebral and pleural ganglia of each side are fused together and are
distinct from the pedal ganglia (Fig. 3E). On the right side, there is a distinct abdominal ganglion, con-
nected to the right pleural ganglion. Optical, rhinophoral, buccal, and esophageal ganglia are also
present and well differentiated. The pedal and parapedeal commissures are enveloped together by
connective tissue with the visceral loop. There are three cerebral nerves leading from each cerebral
ganglion and three pedal nerves leading from each pedal ganglion. From the pleural ganglia lead two
nerves on the right one and three nerves on the left one.

Circulatory and excretory systems. There is a large heart that connects to a single, ramified
blood gland by the aorta. The blood gland is situated covering the central nervous system. The syrinx
is small and pyriform.

GEOGRAPHIC RANGE. — This species was previously known from Puerto Penasco, Sonora,
Mexico, which is situated in the north part of the Mar de Cortés (Sea of Cortez). The present paper
provides the second record of this species, from the La Paz area, Baja California Sur, and Mazatlan,
Sinaloa, and constitutes a range extension of approximately 900 km (Fig. 5). So far this species ap-
pears to be endemic to the Gulf of California.

DISCUSSION

Inuda luarna was originally described from Puerto Penasco, Sonora, Mexico, based on two pre-
served specimens collected by P. Pickens. The original description of this genus and species (Marcus
and Marcus 1967), includes anatomical descriptions and information on the external coloration of the
preserved specimens. According to these authors, /nuda luarna is a whitish species mottled with
brown, due to numerous dots of dark brown pigment. The rhinophores are brown and the gill is darker
than the notum. This coloration is very similar to our specimens from Baja California Sur, which are
brown with creamy white areas and dark brown spots on the center for the dorsum. In addition, the gill
and rhinophores of our specimens are dark brown. Anatomically, the radula described for the Sonora
specimens is identical to that of the specimens from Baja California Sur. However, there are some an-
atomical differences in the reproductive system between the specimens examined by Marcus and
Marcus (1967) and our own material. Our specimens have a smaller prostate and a larger ampulla than
the animal illustrated by Marcus and Marcus (1967, fig. 43). These differences could be due to a dif-
ferent state of maturity in the specimens studied or to intraspecific variation. Despite these differ-
ences, there is no doubt that all the animals belong to the same species.

Marcus and Marcus (1967) considered that the genus /Jnuda resembled Cadlina Bergh, 1879, but
according to these authors the well-developed prostate of Jnuda “makes it impossible to allocate these
genera in the same subfamily.” Thus they erected the new subfamily Inudinae (family
Chromodorididae) to accomodate /nuda luarna. The subsequent references to this genus and species

VALDES AND ANGULLO CAMPILLO: REDESCRIPTION OF CADLINA LUARNA 83

FiGURE 4. Scanning electron micrographs of Cadlina luarna (CASIZ 121104). A. Innermost radular teeth, scale
bar = 43 pm; B. Mid-lateral radular teeth, scale bar = 38 pm; C. Outermost radular teeth, scale bar = 30 um. D. Jaw elements,
scale bar = 23.1 ym.

by Keen (1971), Abbott (1974), Skoglund (1991) and Gonzalez (1993) are based on the paper by
Marcus and Marcus (1967) and not on newly-collected animals. Farmer (1980) re-examined and illus-
trated the type material of this species but did not provide additional anatomical information. Angulo
Campillo (2000) collected additional specimens of this species, which are also re-examined here, and
provided the first data on the external coloration of the living animals. All these authors agreed to
maintain the genus /nuda as valid.

84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No.7

The presence of mantle
glands in /nuda luarna clearly
places this species in the family
Chromodorididae (see Rudman
1984; Gosliner and Johnson
1994; Gosliner and Johnson
1999). These glands were over-
looked by Marcus and Marcus
(1967), who worked with pre-
served material. Of all members
of Chromodorididae, nuda most
resembles Cadlina. Rudman
(1984) reviewed the genus
Cadlina, which according to
him is characterized by having
a heavily spiculose mantle with
small tubercles. The mantle
glands are large, forming a
submarginal row around the
mantle. The branchial leaves
are simple with a tendency to
bipinnate and __ tripinnate
branching. The radula is nar-
row with the number of teeth in the half-row being approximately equal to one-third the number of
rows of teeth. The denticles on the teeth are normally few and large. There is a rachidian tooth with no
central cusp, but on each side of the midline there are one or two large denticles. Most species have a
penis armed with minute hooks. /nuda luarna fits this diagnosis, except for the presence of a large,
central cusp in the rachidian teeth. In our opinion, this small difference does not justify the mainte-
nance of a different genus, and therefore /nuda is here synonymized with Cadlina.

Cadlina luarna is clearly different from other species of the genus. The main diagnostic feature is
the presence of a well-developed prostate, a character used by Marcus and Marcus (1967) to place this
species in a different genus and subfamily. Other species of Cadlina present on the west coast of the
Americas are Cadlina flavomaculata MacFarland, 1905, Cadlina sparsa (Odhner, 1921), Cadlina
linbaughorum Lance, 1962, Cadlina luteomarginata MacFarland, 1966, and Cadlina modesta
MacFarland, 1966. All these species clearly differ from Cadlina luarna in having a white or pale
creamy white background body color (see Behrens 1991).

Gosliner and Johnson (1994) hypothesized that Actinocyclidae is the sister clade to the
Chromodorididae. Rudman (1984) recognized Cadlina to be basal within the Chromodorididae, be-
ing the sister group to a clade containing the genera Glossodoris Ehrenberg, 1831, Verconia
Pruvot-Fol, 1931 and Ardeadoris Rudman, 1984. Recently Gosliner and Johnson (1999) provided a
parsimony-based phylogeny of the Chromodorididae and found Cadlina to be the most basal member
ofaclade containing 7yrinna Bergh, 1898 and Cadlinella Thiele, 1931, which is the sister group to the
rest of the Chromodorididae. The genus Cadlina possesses several plesiomorphic features such as
spiculose body, rachidian teeth, serial seminal receptacle (Gosliner and Johnson 1999). In addition, C.
/uarna retains plesiomorphies present in Actinocyclus, but absent in other members of Cad/ina. These
include a rounded body with an elevated dorsal hump and a well-developed prostate. The arrangement
and size of the mantle glands of C. /uarna also appear to be plesiomorphic. They are smaller than in
most members of Cadlina, and disorganized, whereas in other species of Cadlina they are arranged in
a single submarginal row. Another possible plesiomorphy of C. /uarna is the presence of a large, cen-

Pacific Ocean

FIGURE 5. Geographic range of Cadlina luarna in Mexico.

VALDES AND ANGULLO CAMPILLO: REDESCRIPTION OF CADLINA LUARNA 85

tral cusp in the rachidian teeth. According to this particular external morphology and anatomy,
Cadlina luarna appears to be a very basal member of Cadlina, which is also basal within the
Chromodorididae. Therefore, this species is extremely important to the study of the phylogenetic rela-
tionships within the Chromodorididae and the basal clades of Cryptobranchia.

ACKNOWLEDGMENTS

The authors would like to recognize the generous support of several individuals. Hans Bertsch,
Liza Gomez, Enrique Gonzalez, David Siqueiros, Jose Luis Arreola, Patrick McDonough and the
staff of Proyecto Fauna Arrecifal (Universidad Autonoma de Baja California Sur), provided invalu-
able assistance during the field work. In addition, Hans Bertsch shared with us unpublished informa-
tion on the biogeography and ecology of the opisthobranchs from Baja California that was critical for
the completion of the manuscript.

This paper has been supported by the National Science Foundation through the PEET grant
DEB-9978155, “Phylogenetic systematics of dorid nudibranchs,” to Terrence M. Gosliner of the Cal-
ifornia Academy of Sciences.

LITERATURE CITED

ABBOTT, R. T. 1974. American Seashells. The marine Mollusca of the Atlantic and Pacific coasts of North Amer-
ica, 2nd ed. Van Nostrand Reinhold, New York. 663 pp. + 24 pls.

ANGULO CAMPILLO, O. J. 2000. Moluscos opistobranquios (Mollusca: Opisthobranchiata) de Baja California
Sur, México. B.A. Dissertation, Universidad Autonoma de Baja California Sur, La Paz. 176 pp.

BEHRENS, D. W. 1991. Pacific coast nudibranchs. A guide to the opisthobranchs Alaska to Baja California. Sea
Challengers, Monterey, California. 107 pp.

FARMER, W. M. 1980. Sea-slug gastropods. Farmer Enterprises, Tempe, Arizona. 177 pp.

GONZALEZ, N. E. 1993. Moluscos endémicos del Pacifico de México. Pp. 223-252 in Biodiversidad marina y
costera de México, S. I. Salazar-Vallejo and N. E. Gonzalez, eds. Comision Nacional de Biodiversidad y
CIQRO, México D. F. 865 pp.

GOSLINER, T. M. ANDR. F. JOHNSON. 1999. Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with
a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species.
Zoological Journal of the Linnean Society 125:1—114.

GOSLINER, T. M. AND S. JOHNSON. 1994. Review of the genus Hallaxa (Nudibranchia: Actinocyclidae) with de-
scriptions of nine new species. The Veliger 37:155—191.

ICZN, 1967. Opinion 812. Cadlina Bergh, 1878 (Gastropoda): validate under the plenary powers. Bulletin of
Zoological Nomenclature 24:91—92.

KEEN, A. M. 1971. Sea shells of tropical West America. Marine mollusks from Baja California to Peru, 2nd ed.
Standford University Press, Stanford, California. 1064 pp. + 22 pls.

MARCUS, EVELINE AND ERNEST MARCUS. 1967. American opisthobranch mollusks. Studies in Tropical Ocean-
ography 6:1—256, pl. 1.

RUDMAN, W. B. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: a review of
the genera. Zoological Journal of the Linnean Society 81:115—273.

SKOGLUND, C. 1991. Additions to the Panamic Province Opisthobranchia (Mollusca) literature 1971 to 1990.
The Festivus 22 (Supplement 1):1—27.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park 7
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 8, pp. 87—96, 6 figs., 1 table. October 18, 2000

A New Genus and Species of Soft Coral
(Octocorallia: Alcyoniidae) from South Africa

by

Gary C. Williams Ut | A °
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118
Woods Hole, jy
— bv, 4254"
John Starmer ae oe
Coral Reef Research Foundation,
P.O. Box 1765, Koror, Palau 96940

and

A new genus and species of recently discovered soft coral is described from the shal-
low-water temperate reefs of the eastern Cape Province of South Africa. The new taxon is
somewhat similar to the southern African endemic soft coral Malacacanthus capensis
(Hickson, 1900). However, the new species has sclerites and appears to be monomorphic,
while Malacacanthus capensis does not have sclerites and has dimorphic polyps.

In the past twelve years, several new endemic genera of southern African octocorals have been
described. Included here are the soft coral genera Pieterfaurea Verseveldt and Bayer (1988),
Verseveldtia Williams (1990), Leptophyton Ofwegen and Schleyer (1997), Lampophyton Williams
(2000), and Dimorphophyton Williams (2000); and the pennatulacean genus Amphibelemnon
Lopez-Gonzalez, Gili, and Williams (2000).

This paper reports the discovery and description of an additional new genus and species of soft
coral from the South Coast of South Africa.

METHODS

An examination of recently collected material was made for this study. The material was col-
lected by SCUBA and preserved in 70% ethanol. Sclerites were isolated using sodium hypochlorite.
Micrographs were made using a Kodak MDS100 digital video camera and a Wild M400
photomicroscope. Scanning electron micrographs were made on a Hitachi S-510 scanning electron
microscope. Abbreviations used in the text are as follows: CAS (California Academy of Sciences, San
Francisco), CRRF (Coral Reef Research Foundation, Palau).

A small specimen of Malacacanthus capensis (Hickson, 1900), collected from the same locality
as the new taxon, was examined for comparative purposes. Collection data for this specimen is as fol-
lows: CAS 118498, (Sta. No. SAFR 365), Republic of South Africa, Cape Province, off Port Eliza-
beth, Algoa Bay, White Sands 6, 20 m depth, 26 February 1999, collected by John Starmer with aid of
SCUBA, one whole specimen 15 mm in length.

87

88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 8

SYSTEMATIC ACCOUNT

Family Alcyontidae Lamouroux, 1812

Lanthanocephalus gen. nov.

TYPE SPECIES. — Lanthanocephalus clandestinus sp. nov. by original designation and
monotypy.

DIAGNOSIS. — Growth form upright, unbranched, cylindrical to clavate. Polypary retractile into
distal region of stalk. Polyps monomorphic and retractile, without calyces. Surface coenenchymal
sclerites: small spindles and radiates.

ETYMOLOGY. — The new generic name is derived from the Greek, /anthano (to escape notice;
unknown or unseen) and kephale (a head); in reference to the ability of the polypary to retract into the
distal portion of the stalk.

Lanthanocephalus clandestinus sp. nov.
Figs. 1-6

TYPE MATERIAL. — Holotype: CAS 118500 (Sta. No. SAFR 365), Republic of South Africa,
Cape Province, off Port Elizabeth, Algoa Bay, White Sands 6, on exposed rock, 20 m depth, 26 Febru-
ary 1999, collected by John Starmer with aid of SCUBA, one whole specimen 16 mm in length.
Paratype: CAS 118499, same data as holotype, one 22 mm long specimen cut longitudinally into two
halves.

DIAGNOSIS. — Alcyontid soft corals with upright, unbranched, cylindrical to clavate growth
form. Stalk rigid with rough surface texture. Polyps retractile, without calyces, presumably
monomorphic, restricted to distal portion of the soft coral. Polypary retractile into distal portion of
stalk. Retracted polypary conical to mammiform. Polypary sclerites: small spindles with pronounced
conical tubercles 0.1 1—0.18 mm long. Stalk sclerites: radiates 0.04—0.09 mm in length. All sclerites
colorless. Wet-preserved color dark brown.

DESCRIPTION OF THE HOLOTYPE. — Growth form and size. The holotype is 16 mm in length
and 6 mm in width at its widest portion. The growth form is unbranched and cylindrical to clavate
(Figs. 16832):

Stalk and Polypary. The stalk is relatively rigid and exhibits a tough, somewhat crenulated sur-
face texture. The polypary is retractile into the distal region of the stalk, forming a conical or
mammiform distal terminus (Figs. 1A, B; 2A—C). The retracted polypary comprises 5—12% of the to-
tal body length of the paratype and holotype (Fig. 1A, B).

Polyps. A detailed study of the polyps is not possible since the polypary is almost completely re-
tracted into the distal region of the stalk. However, examination of the tip of the polypary of the
holotype, as well as the interior of the longitudinally sectioned paratype, shows that the polyps are
fully retractile (without calyces) and presumably monomorphic, since no siphonozooids are visible.
Microscopic examination of the longitudinally sectioned paratype as well as a transverse section
through the retracted polypary did not reveal any evidence for the presence of siphonozooids.

Sclerites. The surface coenenchyme of the distal tip of the polypary contains short robust spin-
dles with pronounced, mostly conical tubercles (Fig. 3B). They vary in length from 0.1 1—0.18 mm.
Also present are some radiates. The sclerites of the surface coenenchyme of the stalk are radiates:
mostly eight-radiates, and irregularly-shaped forms that are presumably derived from radiates (Fig.
3A, 4, 5). They vary in length from 0.04—0.09 mm. The interior of the polypary and stalk appears not
to contain sclerites or at most they are very sparse. All sclerites are colorless.

Color of wet-preserved material. Exterior dark chocolate brown throughout; interior uniform
reddish brown.

WILLIAMS AND STARMER: SOUTH AFRICAN SOFT CORAL 89

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FiGure 1. A—B. Lanthanocephalus clandestinus gen. and sp. nov. A. Paratype (CAS 118499); 22 mm in length. B. Holotype
(CAS 118500); 16 mm in length. C-D. Malacacanthus capensis (Hickson, 1900). C. Specimen 80 mm long with re-
tracted polyps, showing capitulate growth form. D. Specimen 75 mm long with retracted polypary. C and D after Williams
(1987:1339, fig. 2).

DISTRIBUTION. — Algoa Bay, eastern Cape Province, South Africa (Fig. 6): 20 m in depth.

ETYMOLOGY. — The specific epithet of the new species is derived from the Latin, clandestinus
(hidden or secret); in reference to the retractile polyps, and the ability of the polypary to withdraw into
the distal portion of the stalk.

REMARKS. — The new taxon is at present known only from the holotype and paratype. Both of
these specimens have the polypary tightly retracted into the distal region of the stalk. It is therefore not
possible to provide detailed descriptions of the polypary or polyps.

DISCUSSION
Lanthanocephalus clandestinus gen. and sp. nov. is sympatric with, and most closely resembles,

the southern African endemic soft coral Malacacanthus capensis (Hickson, 1900). The latter species
has been illustrated or described, or at least mentioned, in several publications under various

90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGURE 2. Lanthanocephalus clandestinus gen. and sp. nov. A. Micrograph of distal portion of wet-preserved paratype with
retracted polypary; length of portion shown = 14 mm. B. Micrograph of distal tip of wet-preserved paratype; length of portion
shown =5 mm. C. Micrograph of distal portion of wet-preserved holotype with retracted polypary; length of portion

shown = 14 mm. D. Malacacanthus capensis (Hickson, 1900). Micrograph of distal portion of wet-preserved specimen show-
ing partly expanded polyps; length of portion shown = 10 mm.

WILLIAMS AND STARMER: SOUTH AFRICAN SOFT CORAL 9]

FIGURE 3. Lanthanocephalus clandestinus gen. and sp. nov. Variation in sclerites from the surface coenenchyme of the
holotype. A. Sclerites from the stalk. B. Sclerites from the distal tip of the polypary. Scale bar for both A and B = 0.1 mm.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 8

FiGuRE 4. Lanthanocephalus clandestinus gen. and sp. nov. Scanning electron micrographs of sclerites from the surface
coenenchyme of the holotype. Lengths: A. 0.07 mm. B. 0.09 mm. C. 0.07 mm. D. 0.08 mm. E. 0.09 mm. F. 0.08 mm G.
08 mm. H. 0.07 mm. I. 0.09 mm.

WILLIAMS AND STARMER: SOUTH AFRICAN SOFT CORAL 95

FIGURE 5. Lanthanocephalus clandestinus gen. and sp. nov. Scanning electron micrographs of sclerites from the surface
coenenchyme of the holotype. Lengths: A. 0.06 mm. B. 0.08 mm. C. 0.08 mm. D. 0.09 mm.

binomens. These names include Heteroxenia capensis, Xenia uniserta, Malacacanthus rufus,
Heteroxenia uniserta, and Malacacanthus capensis, as well as Malacacanthus, and the vernacular
“sunburst soft coral.” The relevant publications include Bayer (1981), Branch and Branch (1981),
Branch et al. (1994), Broch (1939), Day et al. (1970), Hickson (1900, 1931), Kiikenthal (1906), J. S.

94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Indian
Ocean

FIGURE 6. Map of southern Africa showing distributions of Lanthanocephalus clandestinus gen. and sp. nov. (@), and
Malacacanthus capensis (Hickson, 1900) (@). Arrows represent type localities.

Thomson (1910, 1921, 1924), Tixier-Durivault (1954), Williams (1987, 1990, 1992a, 1992b), and
Williams and Alderslade (1999).

Lanthanocephalus clandestinus and Malacacanthus capensis share one important morphologi-
cal character besides having sympatric distributions (Fig. 6; Table 1). Both taxa have polyparies capa-
ble of retraction into the stalk (Figs. 1D, 2B). However, they differ in three significant characters.
Lanthanocephalus clandestinus has sclerites, does not have a cuticle surrounding the stalk, and ap-
pears to be monomorphic since siphonozooids are not evident in the two retracted specimens. The
capitate Malacacanthus capensis (Fig. 1C), on the other hand, does not have sclerites, a reddish or-
ange cuticle covers the epidermis of the stalk, and it is conspicuously dimorphic. In light of these ob-
servations, it is therefore justified to differentiate the two taxa into separate genera. A 15-mm,
wet-preserved specimen of Malacacanthus capensis (CAS 118498) was examined for comparison.
The autozooids are partly expanded and measure 3-4 mm in length (Fig. 2D). Minute siphonozooids
are apparent between the autozooids, each approximately 0.3 mm in diameter.

Ceratocaulon wandeli Jungersen, 1898, from subarctic waters, also has a distinct horny cuticle
surrounding the stalk as in Malacacanthus capensis, but is monomorphic and cylindrical in shape as is
Lanthanocephalus clandestinus. It differs from these two taxa however, in having sclerites that re-
semble minute ovals or rounded platelets.

Species of the paralcyoniid genus Studeriotes Thomson and Simpson, 1909, can also withdraw
the polyparium into the stalk and are also monomorphic, but they differ in having sclerites that are pre-
dominantly large robust spindles, numerous fingerlike branches that compose the polyparium, and
polyps with conspicuous calyces comprised of abundant sclerites. Pertaining to Studeriotes mirabilis
(J. A. Thomson, 1908) from the Andaman Islands, J. A. Thomson and Simpson (1909:8, figs. 3-4)

WILLIAMS AND STARMER: SOUTH AFRICAN SOFT CORAL 95

TABLE |. Comparison of Lanthanocephalus clandestinus and Malacacanthus capensis.

Lanthanocephalus clandestinus Malacacanthus capensis
Maximum length 22 mm 80 mm
Growth form cylindrical to clavate cylindrical to capitate
Sclerites radiates < 0.09 mm, colorless; absent
spindles < 0.18, colorless
Distribution Port Elizabeth Cape Town to East London
Depth range 20m 10-93 m
Polyps presumably monomorphic dimorphic
Stalk cuticle absent present
Polypary shape conical to mammiform usually capitate
(wet preserved)
Polypary retraction into yes yes
stalk
Wet-preserved color dark brown orange polyps, reddish brown stalk

clearly illustrate the fully retracted polyparium within the trunklike stalk, as well as the conspicuous
calyces of the retracted polyps.

ACKNOWLEDGMENTS

Field research in South Africa was made possible by a contract from the United States National
Cancer Institute to the Coral Reef Research Foundation (Palau), and a research permit to Rhodes Uni-
versity Department of Chemistry. We thank Sea Fisheries (South Africa), Dr. Mike Davies-Coleman,
Dr. P. L. Colin and L. J. B. Colin for assistance. We are grateful to Katie Martin (Scientific Publica-
tions, California Academy of Sciences, San Francisco), and Leen van Ofwegen (Nationaal
Natuurhistorisch Museum, Leiden) for their comments and suggestions.

LITERATURE CITED

BAYER, F. M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with
diagnoses of new taxa. Proceedings of the Biological Society of Washington 94:902—947.

BRANCH, G. AND BRANCH, M. 1981. Living Shores of Southern Africa. C. Struik (Pty) Ltd., Cape Town. 272 pp.

BRANCH, G. M., C. L. GRIFFITHS, M. L. BRANCH AND L. E. BECKLEY. 1994. Two oceans—A guide to the marine
life of southern Africa. David Philip, Cape Town and Johannesburg. 360 pp.

BrocH, H. 1939. Some South African shallow water octactinians. Kungliga Fysiografiska sallskapets 1 Lund
forhandlingar 9(6):1—32.

Day, J. H., J. G. FIELD AND M. J. PENRITH. 1970. The benthic fauna and fishes of False Bay, South Africa. Trans-
actions of the Royal Society of South Africa 39(1):1—108.

HICKSON, S. J. 1900. The Alcyonaria and Hydrocorallinae of the Cape of Good Hope. Marine Investigations in
South Africa 1:67—96.

. 1931. The alcyonarian family Xeniidae, with a revision of the genera and species. Great Barrier Reef Ex-
pedition 4(5):137-179.

JUNGERSEN, H. F. E. 1892. Ceratocaulon Wandeli, en ny nordisk Alcyonide. Videnskabelige Meddelelser fra
Dansk Naturhistorisk Forening i Kjobenhavn 1891(1892):234-242.

KUKENTHAL, W. 1906. Alcyonacea. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem
Dampfer ‘Valdivia’ 1898-1899 13(1) Lieferung 1:1—111.

LOPEZ-GONZALEZ, P. J., J.-M. GILI, AND G. C. WILLIAMS. 2000. On some veretillid pennatulaceans from the
eastern Atlantic and western Pacific Oceans (Anthozoa: Octocorallia), with a review of the genus
Cavernularia, and descriptions of new taxa. Journal of Zoology 250(2):201—216.

96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 8

OFWEGEN, L. P. VAN AND M. H. SCHLEYER. 1997. Corals of the South-west Indian Ocean V. Leptophyton
benayahui gen. nov. and spec. nov. (Cnidaria, Alcyonacea) from deep reefs off Durban and on the
KwaZulu-Natal south coast, South Africa. South African Association for Marine Biological Research,
Oceanographic Research Institute, Investigational Report No. 71:1—12.

THOMSON, J. A. 1908. Note on a remarkable alcyonarian Studeria mirabilis, g. et sp. n. Journal of the Royal Mi-
croscopical Society 1908:675—681.

THOMSON, J. A. AND J. J. SIMPSON. 1909. An account of the alcyonarians collected by the Royal Indian Marine
Survey Ship Investigator in the Indian Ocean. II. The Alcyonarians of the littoral sea. Calcutta, The Indian
Museum. 312 pp.

THOMSON, J. S. 1910. The Alcyonaria of the Cape of Good Hope and Natal. Alcyonacea. Transactions of the
Royal Society of Edinburgh 47(3):549-589.

. 1921. South African Alcyonacea. Transactions of the Royal Society of South Africa 9:149-175.

. 1924. Charts and comparisons of the distribution of South African Alcyonaria. With a statement of some
of the problems of their dispersal. Transactions of the Royal Society of South Africa 11:45—84.

TIXIER-DURIVAULT, A. 1954. Les octocoralliaires d’Afrique du sud (I. Aleyonacea). Bulletin du Muséum na-
tional d’Histoire naturelle (2)26(3):385—390.

VERSEVELDT, J. AND F.M. BAYER. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and
Nidaliopsis (Octocorallia: Alcyoniidae and Nidaliidae), with descriptions of two new genera. Zoologische
Verhandelingen 245:1—131.

WILLIAMS, G.C. 1987. The aberrant and monotypic soft coral genus Malacacanthus Thomson, 1910
(Octocorallia: Alcyoniidae) endemic to southern Africa. Journal of Natural History 21:1337—1346.

. 1990. A new genus of dimorphic soft coral from the south-western fringe of the Indo-Pacific

(Octocorallia: Alcyoniidae). Journal of Zoology, London 221:21-35.

. 1992a. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals (Coelenterata,

Anthozoa). Annals of the South African Museum 100(3):249-358.

. 1992b. Biogeography of the octocorallian coelenterate fauna of southern Africa. Biological Journal of

the Linnean Society 46(4):351—401.

. 2000. Two new genera of soft corals (Anthozoa: Alcyontidae) from South Africa, with a discussion of
diversity and endemism in the southern African octocorallian fauna. Proceedings of the California Acad-
emy of Sciences 52(6):65—75.

WILLIAMS, G. C. AND P. ALDERSLADE. 1999. Revisionary systematics of the western Pacific soft coral genus
Minabea (Octocorallia: Alcyontidae), with descriptions of a related new genus and species from the
Indo-Pacific. Proceedings of the California Academy of Sciences 51(7):337-364.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 9, pp. 97-109, 4 figs. October 18, 2000

The Hexandrous Species of Topobea (Melastomataceae)

by

OO Ts
Frank Almeda / c 20 .
Department of Botany, California Academy of Sciences 0 U
Golden Gate Park, San Francisco, California 94115 00s Ho
ee 'G, Ky iA

—

A key and updated geographic and taxonomic notes are presented for the five known spe- ee
cies comprising the hexandrous clade of Topobea. Topobea arboricola, a Panamanian en-
demic described and illustrated here, is distinguished by its six stamens per flower,
2-locular ovary, elliptic to ovate leaves with a caudate-acuminate to long-attenuate apex,
and unappendaged anther connectives. The need for recording field data on taxonomically

useful floral and fruit characters is emphasized.

=~

RESUMEN

Se describe una especie nueva, Topobea arboricola de Panama. Se proveen descripciones,
notas sobre distribucion y fenologia, y discusiones referentes a las afinidades de las cinco
especies de Topobea que tienen seis estambres. También se presenta una clave y mapas de
la distribucion para todas ellas. El color y tamano de los pétalos y estambres, forma de los
poros de las anteras, y el numero de léculos en el ovario son a menudo caracteres
taxonomicos muy utiles en Topobea.

Topobea, with approximately 70 species, is a berry-fruited genus of mostly shrubby epiphytes
and hemiepiphytes restricted to wet forests of tropical America. Over 55 species occur in the
biodiversity hotspot extending from Costa Rica south to Colombia.

Traditionally, Topobea has been defined by its 6-merous isomerous flowers with 12 stamens,
6-locular ovary, and elongate anther thecae with broad confluent apical pores. Although this charac-
terization applies to a majority of species in Topobea, discovery of new species in southern Central
America in the past two decades has necessitated a broadened circumscription of the genus to accom-
modate the accumulation of new and unusual species and a reconsideration of generic limits between
Topobea and its sister genus Blakea (Almeda 1990).

I here provide a review of the most specialized evolutionary line within the genus that I infor-
mally refer to as the hexandrous species. These species, which are largely centered in Panama, share
several synapomorphic characters that include: (1) 6-merous flowers with six stamens, each of which
is attached to the hypanthial torus opposite a calyx lobe; (2) subsessile or short-pedunculate flowers;
and (3) 2-locular or 4-locular completely inferior ovaries. Until recently, most of the species treated
here were known from few collections. A study of additional material generated by my recent field
work and that of colleagues now makes it possible to emend and augment morphological and distribu-
tional information for described species, propose another new species in this alliance, and provide a
preliminary assessment of interspecific relationships within this well-defined clade.

Understanding intraspecific variation in woody epiphytes has always been a challenge. Many
epiphytic species of Melastomataceae are composed of widely dispersed, low density populations.
This coupled with spotty sampling by collectors and the fragmentary preservation or loss of critical

97

98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 9

floral characters on dried specimens has contributed to the paucity of adequate study material. Petal
shape and color, stamen number, morphology of anther pores, and ovary locule number are essential
characters for the delimitation of species in Topobea. Unfortunately, some of these characters are dif-
ficult to evaluate once specimens have been pressed and dried. Ideally, notes on these characters
should be recorded in the field, if the preservation of flowers and fruits in a liquid medium 1s not possi-
ble.

KEY TO THE HEXANDROUS SPECIES OF TOPOBEA

la. Leaves sessile and cordate-clasping; calyx lobes oblong-ovate; petals white, 8-13 mm long; anthers subulate, 5-7 mm
NOR pie ee ant an seers Aa, nd cree ee RS cae Aig TERE ame, Vee a Ga ala) Reet Soe ee T. cordata
1b. Leaves distinctly petioled, varying from elliptic-ovate to obovate or subrotund; calyx lobes lance-triangular; petals
pink, 4.5-7 mm long; anthers bluntly oblong, 2—2.5 mm long.
2a. Ovary 4-locular, anther pores dorsally inclined at the truncate apex; anther connective elevated dorso-basally into a
blontispuniike appendage cr sees ex ieee bs cayr acltotth dude woo. emery Ae hee ae ee T. crassifolia
2b. Ovary 2-locular, anther pores somewhat ventrally inclined at the truncate apex; anther connective unappendaged
dorso-basally (if connective is appendaged then uppermost cauline nodes beset with spreading tufts of hairs).
3a. Leaves thick and leathery when dry, the apex obtuse to rounded, the abaxial surface glandular punctate.
4a. Hemiepiphytic shrub with adventitious roots on the cauline internodes; uppermost cauline nodes beset with
caducous tufts of spreading hairs; leaves ovate-elliptic to subrotund; Cerro Jefe, Panama . . . T. hexandra
4b. Terrestrial shrub without adventitious roots on the cauline internodes; uppermost cauline nodes lacking
tufts of spreading hairs; leaves obovate to oblanceolate; Cerro Tute, Panama......... T. caliginosa
3b. Leaves not thick and leathery when dry, the apex caudate-acuminate, the abaxial surface not glandular
PURI ALE eis tate Ge sy de he er ngeoe Ge Rae oni KD we eer MSI fp ane ee T. arboricola

SPECIES DESCRIPTION

1. Topobea arboricola Almeda, sp. nov.
Fig. |

TYPE. — PANAMA. Bocas del Toro/Chiriqui border: windswept cloud forest on slopes and val-
leys of the Cerro Colorado region, elev. 1450 m, 27 Jan. 1989, Almeda et al. 6456 (holotype: CAS!;
isotypes: MO!, PMA!).

Frutex epiphyticus. Lamina 3.8—7.1 x 1.6—3.9 cm elliptica vel ovata apice caudato-acuminata
vel attenuata basi acuta vel rotundata, 3—5-nervata papyracea et integra, nervis secundariis
nervulisque invisis; petioli 0.9-2.8 cm longi. Flores 6-meri sessiles vel subsessiles in quoque nodo
superiori singuli; bracteae omnino liberae; bracteae exteriores 2.54 x 1.5—-4 mm, ellipticae vel
ovato-ellipticae apice acuto vel obtuso; bracteae interiores 2—3 x 2—3 mm, ovatae vel suborbiculares
apice rotundato. Calycis tubus 0.5 mm longus, lobis 2 mm longis. Petala 5 x 3.5 mm obovata vel
obovato-elliptica apice obtuso. Antherae 6, ca. 2 x 0.5 mm oblongae inter se non cohaerentes;
connectivum nec prolongatum nec appendiculatum. Ovarium 2-loculare et omnino inferum apice
glabro (cono et collo non evoluto).

Epiphytic shrub to 1.5 m tall. Uppermost branchlets glabrous, somewhat compressed and bluntly
two-edged; older branches becoming + rounded with nodular leaf scars. Mature leaves of a pair equal
to somewhat unequal in size, glabrous throughout; petioles 0.9-2.8 cm long; mature blades
papyraceous, 3.8—7.1 cm long and 1.6—3.9 cm wide, elliptic to ovate, apex caudate-acuminate to at-
tenuate, base acute varying to rounded, margin entire, 3—5-nerved, the outermost intramarginal pair
often depressed and inconspicuous, the transverse secondary veins typically not elevated or conspicu-
ous. Flowers erect, solitary in leaf axils of uppermost branches, sessile, subsessile, or with short
(1-2 mm) ill-defined peduncles formed by the compressed bases of the outer floral bracts. Floral

ALMEDA: TOPOBEA 99

FiGURE |. Topobea arboricola Almeda. A. habit, x ca. '4; B. representative leaf (abaxial surface), x 1; C. young fruiting
hypanthium with decussate floral bracts, x 6; D. stamens, ventral view (left) and partial profile view (right), x 7; E. petal
(adaxial surface), x 7; F. berry (top view) showing calyx lobes, ovary summit, and torus with staminal filament scars, x 4; G.
seeds, x 15. (A, B from the holotype; C-E from McPherson 12813, MO; F, G from McPherson 7739, CAS.)

100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 9

bracts free and entire, glabrate or sparingly beset abaxially (mostly toward the apex) with a lanate or
stellulate-lepidote, coppery brown indument; outer bracts 2.5—4 x 1.5—4 mm, concave, elliptic to el-
liptic-ovate, apex acute to obtuse; inner bracts 2—3 x 2-3 mm, depressed-ovate to semicircular, apex
rounded. Calyx tube 0.5 mm long; calyx (in fruit) erect, 2 mm long and 2 mm wide basally, deltoid to
deltoid-ovate, entire but callose-thickened along the interlobe sinuses, sometimes sparingly beset at
the abaxial apex with an indument like that of the floral bracts. Petals 6, sparingly beset with
disc-shaped glands when dry, 5 x 3.5 mm, translucent pink, obovate to elliptic-obovate, apex obtuse,
entire. Stamens 6, filaments complanate and glabrous, 2.5—-3 mm long; anthers 2 mm long, 0.5 mm
wide, pale yellow, oblong with two broad pores at the truncate apex; connective barely thickened and
unappendaged. Ovary completely inferior, 2-locular, apex glabrous, stylar scar evident but not ele-
vated into a prominent cone or stylar collar. Style somewhat declinate and sigmoid apically, glabrous,
5 mm long; stigma punctiform. Berry 4—6 =< 4.5—-6 mm, white at maturity. Seeds cuneate to narrowly
deltoid, | mm long, tan with a smooth, glossy testa.

PHENOLOGY. — Flowering in February, May, July and August; fruiting specimens have been
collected in July, December, and January.

DISTRIBUTION. — Known only from wet cloud forests of western Panama from Valle de la
Sierpe (Chiriqui) east to Cerro Tute (Veraguas) at 1000-1500 m (Fig. 2).

PARATYPES. — PANAMA. Bocas del Toro/Chiriqui border: above Fortuna dam, along divide
trail, ca. 8°45'N, 82°15'W, 4 Dec. 1985, McPherson 7739 (CAS, MO). Chiriqui: Fortuna dam area,
along Quebrada Bonita to E of road, 8°45'N, 82°13'W, 8 Feb. 1984, Churchill et al. 4759 (MO); Valle
de la Sierpe, en direccion SE a lo largo de Quebrada Bonita, 17 May 1987, Correa et al. 5092 (MO,
PMA); Fortuna dam area, N of reservoir, ridge along continental divide and southward from
Quebrada de Arena, Aug. 1984 (no day given), D’Arcy & Todzia 15959 (CAS, MO). Veraguas: trail to
Reserva Bioldgica Serrania de Tute about 0.7 km beyond the Escuela Agricola Rio Piedra just outside
of Santa Fé, 18 Feb. 1996, Almeda et al. 7622 (CAS, MO, PMA, US); Distrito de Santa Fé, Serrania de
Tute, 8°33'N, 81°07'W, 5 Jul. 1996, Aranda et al. 2676 (CAS, SCZ); Distrito de Santa Fé, Serrania de
Tute, 8°33'N, 81°07'W, 5 Jul. 1996, Aranda et al. 2657 (CAS, SCZ); vicinity of Cerro Arizona-Tute,
above Santa Fé and Altos Piedra, along trail to summit, 8°30'N, 81°10'W, 28 Jul. 1988, McPherson
12813 (MO).

DISCUSSION. — Topobea arboricola appears to be a true epiphytic shrub that never has root con-
tact with the ground. It is distinguished by its 2-locular ovary, comparatively long petioles
(0.9-2.8 cm), elliptic to ovate leaves with a caudate-acuminate to long-attenuate apex, and short ob-
long anthers with unappendaged connectives.

Of the three hexandrous species of Topobea with 2-locular ovaries, T. arboricola is most similar
to T. caliginosa. The latter differs in being a terrestrial shrub and in having obovate leaves with an ob-
tuse to rounded apex, shorter petioles (0.5—1 cm), longer floral peduncles (3—5 mm), and abaxial fo-
liar surfaces that are inconspicuously glandular-punctate.

All collections of 7. arboricola come from a scattering of localities along the Cordillera Central
of Panama. For an epiphyte with a dispersed population structure, this species 1s remarkably uniform
in vegetative and reproductive characters. An exception to this homogeneity is exhibited by the Cerro
Tute population which has uniformly ovate leaves (vs. elliptic leaves).

ETYMOLOGY. — The epithet for this species, arboricola, is derived from the Latin word, arbor,
meaning dwelling in a tree, in reference to its epiphytic habit in the forest canopy.

101]

ALMEDA: TOPOBEA

BJOOUOdIe "| @

‘euleueg Ul Mj/0I140G4D vaqgodo] Jo uoNNqiysiq ‘7 AUNDIA

102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 9

2. Topobea caliginosa Almeda, Proc. Calif. Acad. Sci. 46(14):317. 1990.

Blakea micrantha Almeda, Rhodora 82:614. 1980.

TYPE. — PANAMA. Veraguas: Cerro Tute ca. 10 km NW of Santa Fé on ridgetop in cloud forest
above 1000 m, 19 June 1975, Mori 6765 (holotype: CAS!; isotype: MO!).

Rigidly branched terrestrial shrub to | m tall. Upper cauline internodes quadrangular to distinctly
carinate. Young vegetative buds and juvenile foliage commonly beset with a sparse coppery brown
furfuraceous indument, otherwise glabrous throughout. Leaves coriaceous and glabrous when dry but
inconspicuously punctate abaxially, 14 x 0.4-1.6 cm, obovate to oblanceolate, apex obtuse to
rounded, base acute to attenuate, the margin entire, often revolute when dry, 3-nerved; petioles
0.5—1 cm long. Flowers solitary in the axils of distal branches, sessile, subsessile or with outermost
floral bracts compressed into an ill-defined peduncle 3-5 mm long. Floral bracts rounded at the apex,
sparsely floccose to glabrous at maturity, margin entire; outer bracts 3-5 x 2—3 mm, fused at the base
for 1-2 mm, elliptic-lanceolate; inner bracts 3-4 x 2-3 mm, free, elliptic-ovate. Calyx lobes bluntly
deltoid, 1.5 mm long and 2 mm wide basally. Petals 6, glabrous but sparsely verrucose abaxially when
dry, 4.5—6 x 2.5-3 mm, pink, ovate to elliptic-ovate, apex obtuse to rounded, entire. Stamens 6, fila-
ments 2—3.5 x 0.5 mm, erect, glabrous; anthers free, 2 x 0.5—1 mm, linear-oblong and erect, each
with two confluent pores at the truncate apex; connective not thickened or appendaged dorsally.
Ovary completely inferior, 2-locular, glabrous at the apex which is elevated into a short cone. Style
glabrous, 5-6 mm long; stigma punctiform. Berry 4-5 « 3-5 mm. Seeds mostly | mm long, cuneate
to narrowly pyriform.

PHENOLOGY. — Flowering specimens were collected in February, April, and June; fruiting spec-
imens collected in April, June, and July.

DISTRIBUTION. — Western Panama where it is known only from Cerro Tute (Veraguas) at
1400-1453 m (Fig. 3).

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Veraguas: windswept summit of
Cerro Tute, 18 Feb. 1996, Almeda et al. 7627 (CAS, MO, PMA); distrito de Santa Fé, Serrania de
Tute, 8°33'N, 81°07'W, 5 Jul. 1996, Aranda et al. 2724 (CAS, SCZ); summit of Cerro Tute above
Escuela Agricola Alto de Piedra, just W of Santa Fé, 8°32'N, 81°07'W, 5 June 1982, Knapp &
Dressler 5394 (CAS, MO).

DISCUSSION. — This species was originally described as an epiphytic shrub based on informa-
tion provided by the collector of the type and only known collection at the time (Almeda 1980). Dur-
ing recent field work I have found 7. caliginosa growing only as a terrestrial shrub above tree line on
the shrubby summit of Cerro Tute.

Three other congeners in this hexandrous group of species also occur on Cerro Tute—T.
arboricola, T. cordata, and what may be an undescribed taxon most closely related to 7. hexandra.
The two former taxa appear to be true epiphytic shrubs that grow on trees within the cloud forest zone.
The third entity, which grows in the upper zone just below tree line, is a vinelike secondary
hemiepiphyte that germinates terrestrially, ascends nearby trees by adventitious roots, and later be-
comes epiphytic by losing root contact with the ground. This sympatry and parapatry suggest that
strong isolating mechanisms are operating to maintain phenotypic distinctions among four closely re-
lated species. That this all occurs on a tropical mountain of 1453 m elevation makes it all the more re-
markable.

103

ALMEDA: TOPOBEA

‘eueueY Ul DApUuDxaYy ‘J, pue “DIDp10I “J “DSou1s1]D2 vagodo] Jo suoyNqiysiq ‘¢ JANDA

BJPUeXeY "| #
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104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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3. Topobea cordata Gleason, Phytologia 3:354. 1950.

TYPE. - PANAMA. Coclé: Cerro Pajita, north of El Valle de Anton, elev. 1000-1200 m, Allen
4178 (holotype: NY!; isotype: MO!)

Epiphytic shrub to | m tall. Uppermost internodes + compressed and quadrisulcate, the angles
distinctly carinate sometimes varying to narrowly winged; older branches + rounded to
rounded-quadrate. Leaves of a pair sessile, equal to slightly unequal in size, the younger ones of a pair
typically fused basally at the node into a narrow collar, glabrous throughout; mature blades
coriaceous, 5.1—-12.9 cm long and 3—5.8 cm wide, cordate-clasping to subcordate or ovate, apex
acute, base rounded to cordate, margin entire, 5—7-nerved, the secondary veins typically not evident.
Flowers erect, coconut-scented (fide McPherson 8738), solitary or paired in each upper leaf axil;
peduncles 34 mm long, conspicuously compressed. Floral bracts entire, glabrate or sparingly beset
abaxially (mostly toward the apex) with a coppery brown lanate indument; outer bracts
0.8—1.1 x 0.6-0.9 cm, fused at the base for 2 mm, elliptic, apex acute to obtuse; inner bracts
0.8—0.9 x 0.4—0.6 cm, free to the base, oblong-ovate, apex rounded. Calyx tube 1—1.5 mm long; calyx
lobes (at anthesis) 4.5 mm long and 3 mm wide, oblong-ovate, + entire, essentially glabrous but some-
times beset with a coppery brown lanate indument toward the apex on both surfaces. Petals 6,
0.8—1.3 x 0.40.7 cm, translucent-white, obovate, apex rounded, entire. Stamens 6, filaments
complanate and glabrous, 6-7 mm long; anthers 5—7 mm long, I—1.5 mm wide, yellow, subulate with
two broad confluent ventrally inclined apical pores; connective not conspicuously thickened and
unappendaged. Ovary completely inferior, 4-locular, apex glabrous and elevated into a blunt basally
swollen cone 2 mm long. Style somewhat declinate and sigmoid apically, glabrous, 8-9 mm long;
stigma truncate. Berry | x | cm, red at maturity. Seeds narrowly pyriform to cuneate, |—1.5 mm long,
beige, the testa smooth and glossy.

PHENOLOGY. — Flowering from February through May, September and probably during inter-
vening months; the only fruiting specimen was collected in September.

DISTRIBUTION. — Wet cloud forests of western Panama from the Fortuna region (Chiriqui) east
to Cerro Pajita (Coclé) at 1050-1250 m (Fig. 3).

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Chiriqui: Fortuna dam region, along
Quebrada Arena near continental divide, 8°45'N, 82°15'W, 9 Mar. 1986, McPherson 8738 (CAS,
MO); Fortuna cauce de Quebrada Arena, 9 Apr. 1987, Valdespino et al. 639 (CAS, PMA). Veraguas:
Cerro Tute, trail past agricultural school near Santa Fé, 17 Sep. 1979, Antonio 1875 (CAS, MO).

DISCUSSION. — The type specimen of this species lacks petals and stamens which accounts for
the incomplete descriptions provided in the protologue (Gleason 1950) and subsequent treatment of
this species in the Flora of Panama (Gleason 1958). The collection of flowering material with attached
stamens facilitated a critical assessment of its generic placement and relationships to other species
with six antesepalous stamens (Almeda 1990).

Topobea cordata is characterized by a distinctive suite of characters including sessile
cordate-clasping leaves, oblong-ovate calyx lobes, comparatively large (8—13 mm) translucent white
petals, and subulate anthers with ventrally inclined apical pores. Topobea cordata has a 4-locular
ovary like 7. crassifolia, but the latter has petiolate leaves, lance-triangular calyx lobes, smaller
(5-7 mm) pink petals, and short (2—2.5 mm) apically truncate anthers.

One enigmatic collection (McPherson 9864, CAS) from the Fortuna region of western Panama
resembles T. cordata in its overall vegetative morphology and white petals. In calyx shape and stami-
nal details, however, it is a good match for T. crassifolia, the species to which T. cordata is most
closely related. In the size of its floral bracts, hypanthia, and petals this collection also approaches 7.
crassifolia. This kind of character sorting is suggestive of hybridization or introgression between T.
cordata and T. crassifolia. Typical T. cordata has also been collected at Fortuna. I have not seen au-

ALMEDA: TOPOBEA 105

thentic material of 7. crassifolia from Fortuna but it may well occur there since its range extends both
east and west of that region.

4. Topobea crassifolia (Almeda) Almeda, Proc. Calif. Acad. Sci. 46(14):318. 1990.

Blakea crassifolia Almeda, Rhodora 82:612. 1980.

TYPE. — PANAMA. Coclé: La Mesa above El Valle in forest on both sides of junction with road
to Cerro Pilon, ca. 800 m, 21 Jul. 1974, Croat 25430 (holotype: CAS!; isotypes: MO!, US!).

Epiphytic shrub to | m tall. Uppermost branches quadrangular to rounded, glabrous throughout.
Leaves thick, succulent or semisucculent, coriaceous or chartaceous when dry,
1.5—6.5(—12) x 14.3 cm, ovate to elliptic-ovate or elliptic-lanceolate, apex acute to attenuate vary-
ing to cuspidate or mucronate, base rounded to cordate, the margin entire, 3—5-nerved, often with only
the median nerve elevated and conspicuous abaxially, essentially glabrous throughout; petioles
0.1—0.4 cm long. Flowers solitary, paired or in subfasciculate clusters of three to five in axils of distal
branches, sessile, subsessile or with ill-defined peduncles |—2 mm long. Floral bracts glabrous or
sometimes beset with an inconspicuous lanate or floccose indument distally, margin entire; outer
bracts 5—8 x 2.5—4 mm, fused at the base for about 1 mm, elliptic-lanceolate, apex acute to obtuse or
rarely rounded; inner bracts 4-7 x 2.5-4 mm, free but closely subtending the hypanthium, ellip-
tic-ovate, apex acute to rounded or broadly truncate. Calyx lobes lance-triangular, 34.5 mm long and
2—2.5 mm wide between sinuses. Petals 6, glabrous and coarsely verrucose, sometimes fringed with a
scattering of minute matted hairs, 5—7 x 1.5—2.5 mm, reportedly white or pink, oblong-lanceolate to
narrowly oblanceolate, apex acute, base somewhat clawed, entire. Stamens 6, filaments
3.54.5 x 0.5 mm, erect, glabrous; anthers free, 2—2.5 x 0.5 mm, linear-oblong and erect distally,
each with two confluent dorsally inclined apical pores; connective thickened dorso-basally into a
spurlike appendage. Ovary completely inferior, 4-locular and glabrous at the truncate apex. Style gla-
brous, 6 mm long; stigma punctiform. Berry 6 x 5 mm. Seeds mostly | mm long or less, ranging from
ovoid or clavate to lunate or pyriform.

PHENOLOGY. — Flowering in February and May through August; fruiting collections have been
made in May and August.

DISTRIBUTION. — Cloud forests in central Costa Rica from Braulio Carillo National Park (San
José) and vicinity disjunct to Panama from Cerro Colorado (Bocas del Toro) and El Valle de Anton
(Coclé) to El Llano-Carti region (Comarca de San Blas) at 350—1750 m (Fig. 4).

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela/Heredia border: Colonia
Virgen del Socorro along road from Costa Rica #9 to the Colonia, 2 Aug. 1976, J. & K. Utley 5629
(CAS, DUKE, F). San José: Parque Nacional Braulio Carillo. La Montura, 25-30 Jul. 1982, Todzia et
al. 1964 (NY). PANAMA. Bocas del Toro/Chiriqui border: Cerro Colorado, road along top, 13 Aug.
1977, Folsom et al. 4681 (CAS). Coclé: La Mesa, 4 km N of El Valle de Anton, 6 May 1981, Sytsma et
al. 4364 (CAS, MO); along trail to La Mesa about 4.5 miles beyond El Valle de Anton, 21 May 1970,
Wilbur & Luteyn 11697 (CAS, DUKE, F, MO, US). Comarca de San Blas: El Llano-Carti road.
Nusagandi, 19 km from Interamerican Hwy, 9°19’N, 78°55'W, 26 Aug. 1984, de Nevers & de Leon
3767 (CAS, MO); El Llano-Carti road, km 19.1, 9°19'N, 78°55’W, | Jul. 1985, de Nevers 5943 (CAS,
MO); entrada a Nergan Igar, km 15 de la carretera Llano-Carti, 9°20'N, 78°58’W, 2 Jul. 1994,
Galdames et al. 1243 (CAS, SCZ).

DISCUSSION. — Among the hexandrous species, 7. crassifolia is the most widely distributed and
has the broadest elevational amplitude. Like many woody epiphytes, the fragmented but small popu-
lation structure of this species has evidently promoted morphological divergence.

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‘eueueg pure vory eySOD Ul DI/Ofissv49 Dagodo] Jo UONNqINSIG “p ANNI

BI|OJISSEIO ‘Le

ALMEDA: TOPOBEA 107

Most of the known populations of 7. crassifolia can be distinguished by leaf shape alone. Plants
from the type locality near El Valle, Panama, have thick semisucculent leaves that are ovate to ellip-
tic-ovate and bluntly apiculate to obtuse or rounded apically. The Cerro Colorado population in west-
ern Panama has smaller (4.5—5 x 1.7—2.5 cm) elliptic to elliptic-ovate leaves that are coriaceous. The
northernmost population from central Costa Rica has longer elliptic-ovate to elliptic-lanceolate
leaves (5.3—12 cm x 3—3.4 cm) that are thinner, flexuous, and acute to attenuate apically. All of these
populations are otherwise identical in other vegetative and reproductive characters. Because of this, I
here emphasize their unity and underlying similarities by recognizing a single variable species. Varia-
tion of this kind, while striking, is not unprecedented among other epiphytic Melastomataceae.
Leandra subulata Gleason, for example, is another woody epiphytic melastome with many geograph-
ical variants and a comparable montane distribution in Costa Rica and Panama.

Another population from the El Copé region of west-central Panama may also prove to be a geo-
graphical variant of 7. crassifolia. It has ovaries with 4-locules like 7. crassifolia but its thick leathery
leaves are narrowly elliptic, 2.3-3 cm wide, and uniformly long-acuminate basally and apically. This
variant is known from two specimens, Al/meda et al. 7650 (CAS) and Folsom & Robinson 2437
(CAS), both of which are only in bud. These specimens are tentatively excluded from my concept of T.
crassifolia, pending study of additional material.

5. Topobea hexandra Almeda, Proc. Calif. Acad. Sci. 46(14):320. 1990.

TYPE. — PANAMA. Panama: Cerro Jefe, along summit road and along trail into the Chagres
Valley, elev. ca. 900 m, 19 Feb. 1988, Al/meda et al. 5837 (holotype: CAS!; isotypes: CR!, DUKE!, F!,
MO!, NY!, PMA!, TEX!, US!).

Hemiepiphytic shrub to | m tall adhering to the bark of host trees by nodal and internodal adven-
titious roots. Upper cauline internodes quadrate to quadrisulcate, glabrous or sparsely covered with
spreading, caducous, glandular hairs 1-2 mm long, as are the young petioles and both surfaces of ju-
venile leaves; older branches rounded, the leaf scars typically swollen. Uppermost nodes beset with
brown spreading hairs. Vegetative buds copiously covered with a deciduous brown indument of
stellate-lepidote hairs. Leaves coriaceous and glabrous throughout but inconspicuously punctate
abaxially, 2—3.9 x 1.1—3 cm, suborbicular to elliptic-ovate, apex rounded varying to obtuse, base ob-
tuse to rounded, margin entire, 3-nerved, often with an additional inconspicuous intramarginal pair;
petioles 0.5—1.4 cm long. Flowers erect, solitary or paired in the leaf axils of distal branches, sessile or
subsessile with short (to 1 mm) ill-defined peduncles formed by the compressed bases of the outer flo-
ral bracts. Floral bracts thick and semisucculent, free and entire, sparingly stellulate-furfuraceous
abaxially; outer bracts 5—6.5 x 3-5 mm, concave, ovate to elliptic-ovate, apex obtuse to bluntly
mucronate; inner bracts 4—5 x 4-6 mm, broadly ovate to suborbicular, apex rounded. Calyx tube
1 mm long; calyx lobes erect, 2 x 2-2.5 mm, ovate to deltoid-ovate, entire but irregularly roughened
along interlobe sinuses, sparingly stellate-lepidote. Petals 6, liberally covered with hyaline
disc-shaped glands when dry, 6.5—7 x 4 mm, pink, elliptic-obovate, apex obtuse. Stamens 6, fila-
ments 3 mm long and somewhat declinate; anthers free, 2 x 1 mm, pale yellow turning brownish or-
ange with age, oblong, each with two ventrally inclined pores at the broadly rounded apex, connective
slightly thickened and dilated dorso-basally at the filament insertion into a blunt spur up to 0.25 mm
long. Ovary completely inferior, 2-locular, glabrous at the apex and not modified into a cone or collar.
Style glabrous, 5.5 mm long; stigma punctiform. Berry 5S—6 x 4—7.5 mm. Seeds I—1.5 mm long,
beige, bluntly deltoid.

PHENOLOGY. — The only known flowering specimen was collected in February; fruiting speci-
mens have been collected in February, September, October, and December.

108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DISTRIBUTION. — Known only from low cloud forests at the summit of Cerro Jefe east of the Ca-
nal Area in central Panama at 900—1025 m (Fig. 3). The Cerro Jefe region was an important island
refugium from the middle Miocene until the land bridge between North America and South America
was established ca. 3.5—2.4 million years ago (Graham 1985; Lewis 1971). This geologic history is re-
flected in the high incidence of vascular plant endemism on Cerro Jefe. Of the approximately 1230
species of plants thought to be endemic to Panama (Carrasquilla 1997), about 150 occur on Cerro Jefe
including 6 locally endemic species of Melastomataceae.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Panama: summit and S facing slopes
of Cerro Jefe, 9 Feb. 1978, Almeda & Nakai 3459 (CAS); Cerro Jefe along summit road beyond the
Intel tower, 3 Feb. 1996, Almeda et al. 7495 (CAS); Cerro Jefe, along trail on ridge running NE from
summit, 18 Dec. 1974, Mori & Kallunki 3755 (MO); Cerro Jefe, 29 Oct. 1980, Sytsma 2007 (MO);
Cerro Jefe, road leading N from summit, 26 Sep. 1975, J. T. & F. Witherspoon 8552 (MO).

DISCUSSION. — As noted in the protologue, 7. hexandra is acommon hemiepiphyte in the sum-
mit forest of Cerro Jefe, but it appears to be overlooked by most collectors because it has small flowers
that are rarely evident to the casual observer without the aid of binoculars (Almeda 1990). This species
is variable in indument characters. The uppermost cauline internodes can either be glabrous or
sparsely covered with spreading glandular hairs. These glandular hairs are also produced on petioles
of juvenile foliage and on upper and lower surfaces of some young leaves, but they commonly fall
away with age.

Topobea hexandra is readily distinguished from the other hexandrous species with 2-locular ova-
ries by its ovate-elliptic to subrotund leaves, dorso-basally appendiculate anther connectives, and
dark brown tufts of hairs on the uppermost nodes.

What may be an extreme variant or perhaps an undescribed taxon is known from two collections
(Almeda et al. 7621, CAS and Aranda et al. 2733, CAS) made on Cerro Tute, Panama, in 1996. This
entity is also a hemiepiphyte but it has small leaves (0.9—1.4 x 0.4—1 cm) that are obovate with a
rounded to emarginate apex. One of these collections is in bud and the other is in young fruit. In view
of the small sample size and the lack of mature flowers and fruits, no taxonomic disposition of the
Cerro Tute population is possible at this time.

ACKNOWLEDGMENTS

I thank Jenny Speckels for preparing the line drawings in Figure 1; Gerrit Davidse, Gordon
McPherson, Mireya D. Correa, and the Smithsonian Tropical Research Institute for logistical support
in Panama; and the curators and staffs of the following herbaria for special loans and assistance during
study visits: CAS, CR, DUKE, F, INB, MO, NY, PMA, SCZ, and US. Field work was supported, in
part, by U. S. National Science Foundation Grant BSR 8614880 (Flora Mesoamericana), the Nathan
Jay Friedman Fund and the G. Lindsay Field Research Fund of the California Academy of Sciences.

LITERATURE CITED

ALMEDA, F. 1980. Central American novelties in the genus Blakea (Melastomataceae). Rhodora
82(832):609-615.

. 1990. New species and new combinations in Blakea and Topobea (Melastomataceae), with an historical
perspective on generic limits in the tribe Blakeeae. Proc. California Acad. Sci. 46(14):299-326.

CARRASQUILLA, L. G. 1997. Cerro Azul-Cerro Jefe Region, Panama. Pp. 221—225 in Centres of plant diversity: A
guide and strategy for their conservation (Volume 3. The Americas), S. D. Davis, V. H. Heywood, O.
Herrera-MacBryde, J. Villa-Lobos, and A. C. Hamilton, eds. WWF-World Wildlife Fund for Nature and
IUCN-The World Conservation Union. IUCN Publications Unit, Cambridge, U. K.

GLEASON, H. A. 1950. Observations on tropical American melastomes. Phytologia 3:345—360.

ALMEDA: TOPOBEA 109

. 1958. Melastomataceae. /n Flora of Panama, R. E. Woodson, Jr. and R. W. Schery, eds. Ann. Missouri
Bot. Gard. 45:203-304.

GRAHAM, A. 1985. Vegetational paleohistory studies in Panama and adjacent Central America. /n The botany
and natural history of Panama, W. G. D’Arcy and M. D. Correa A., eds. Monogr. Syst. Bot. Missouri Bot.
Gard. 10:161—176.

Lewis, W. H. 1971. High floristic endemism in low cloud forests of Panama. Biotropica 3:78-80.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

\\

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 10, pp. 111—124, 7 figs. October 18, 2000

Two New Species of Chromodorididae (Mollusca: Nudibranchia)
from the Tropical Indo-Pacific, with a Redescription of
Hypselodoris dollfusi (Pruvot-Fol, 1933)

by

Terrence M. Gosliner OG] ie
t

and 2000
David W. Behrens

Department of Invertebrate Zoology and Geology, C eee Ae bop rienasey
Golden Gate Park, San Francisco, California 94118

=“.

ne

Two new species of the family Chromodorididae are described. Chromodoris buchananae
is known from northern New South Wales, Australia. It is similar in appearance to several
members of the Chromodoris quadricolor complex, but differs in its color pattern with a
creamy white body with brown flecks, white longitudinal lines and an orange marginal
band. Hypselodoris dollfusi (Pruvot-Fol, 1933) was originally described from the Gulf of
Suez and is here documented for the first time since its original description. It is also known
from the United Arab Emirates. It is readily distinguished from other members of the ge-
nus by its opaque white body with pink-purple rings and yellow-orange marginal band. It
is similar in its anatomy to several species of the monophyletic clade containing H. fucata,
H. kaname, H. koumacensis and H. paulinae. The color pattern, distribution of mantle
glands and radular tooth morphology clearly separate H. dollfusi from other members of
this clade. Hypselodoris babai is known from Okinawa. It can be distinguished by its
red-brown body color with opaque white markings. It is similar in appearance to H.
bullocki in having an elevated gill sheath, but differs in its coloration and by the presence of
mantle glands.

At the end of the twentieth century, numerous papers described new species of Chromodorididae
from the tropical Indo-Pacific (Rudman 1984, 1986, 1987, 1995; Baba 1995, 1996; Hamatani 1995;
Gosliner and Behrens 1998; Johnson and Gosliner 1998; Gosliner and Johnson 1999; Schrodl 1999;
Valdés et al. 1999). These papers added much to our knowledge of biodiversity of the region and also
clarified higher systematic relationships within the family. Most of these papers have focused on spe-
cies that have been found along the margins of the Pacific and Australasian plates and have added
many new taxa from Japan, Philippines, Papua New Guinea, and Australia. Recently, we were pro-
vided with additional material of two undescribed, large chromodorids from Australia and Okinawa.
This paper describes the anatomy and systematic relationships of these additional taxa and provides a
redescription of Hypselodoris dollfusi from the United Arab Emirates.

11]

112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 10

SPECIES DESCRIPTONS

Family Chromodorididae Bergh, 1891
Genus Chromodoris Alder and Hancock, 1855

Chromodoris buchananae sp. nov.
Figs. 1A, 2A-D, 3A-D

TYPE MATERIAL. — HOLOTYPE: Australian Museum C. 383578, specimen dissected, South
Solitary Island, Coffs Harbour, New South Wales, Australia, 15 m depth, February 1997, Carol Bu-
chanan.

ETYMOLOGY. — Chromodoris buchananae is named for friend and colleague Carol Buchanan
who first discovered this species, and provided us with the holotype.

DISTRIBUTION. — Thus far, this species is known only from the type locality, South Solitary Is-
land, Coffs Harbour, New South Wales, Australia.

EXTERNAL MORPHOLOGY. — The living animal (Fig. 1A) was approximately 48 mm in length.
The body is a translucent cream color with small brown flecks dispersed over the notum. Around the
margin of the notum there is a wide orange band. Within this band is a thin opaque white band. There
are two opaque white lines running from just posterior of the rhinophores to the side of the gills
pocket. There is also a short white line anterior to each rhinophore, and a line forming a posterior to the
gills. The hyponotum and posterior end of the foot are cream colored with a broad orange marginal
band, similar to that on the notum. The posterior end of the foot has a white line forming a “V”, and
there are several white lines of varying length along the side of the hyponotum. The rhinophores and
gill are orange. There are 7 unipinnate branchial leaves forming the branchial plume. The perfoliate
rhinophores bear about 20 lamellae.

MANTLE GLANDS. — The subcutaneous mantle glands (Fig. 2A) form a discontinuous
submarginal band around the notum of the animal. They are absent from the anterior portion of the an-
imal to just behind the rhinophores. These glands consist of highly ramified, spherical lobes that are
clustered together, resembling bunches of grapes (Fig. 2B).

BUCCAL ARMATURE. — The muscular portion of the buccal mass is approximately equal in
length to the oral tube. At the anterior end ot the muscular portion of the buccal mass is a chitinous la-
bial cutical, which bears numerous jaw rodlets. These rodlets (Fig. 3A) are elongate with spreading,
bifid apices. The radular formula is 81 x 63.0.63. There is no trace of a row of rachidian teeth. The in-
nermost lateral teeth (Fig. 3B) are broad and quadrangular. There are three to four elongate denticles
on the inner side of the elongate, triangular primary cusp and four to five denticles on the outer side of
the cusp. The next successive inner lateral teeth lack denticles on the inner side of the cusp, but have
three to four denticles on the outer side. The middle lateral teeth (Fig. 3C) are elongate with 7-9
denticles on the outer side of the broader cusp. The outermost teeth (Fig. 3D) are unevenly curved with
an abbreviated portion containing 3—7 short, rounded denticles below the small primary cusp.

REPRODUCTIVE SYSTEM. — (Fig. 2C) The ampulla is thick and tubular, narrowing somewhat be-
fore bifurcating into an oviduct and vas deferens. The short oviduct enters the female gland mass near
the albumen gland. The prostatic proximal portion of the vas deferens is folded over itself once before
it narrows markedly into the relatively short, muscular, ejaculatory portion. The ejaculatory portion

—

FIGURE |. Living animals. A. Chromodoris buchananae sp. nov., specimen from South Solitary Island, Coffs Harbour, New
South Wales, Australia, photograph by C. Buchanan. B. Hypselodoris dollfusi (Pruvot-Fol, 1933), specimen from Khor
Fakken, United Arab Emirates, photograph by C. Harris and L. Betts. C. Hypselodoris babai sp. nov., specimen from Seragaki,
Okinawa, photograph by R. Bolland.

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE

114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 10

FIGURE 2. Chromodoris buchananae sp. nov. A. Subcutaneous glandular network. B. Mantle glands, enlarged,
scale = 1.0 mm. C. Reproductive system, am = ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland
mass, p = penis, pr = prostate, rs = receptaculum seminis, vg = vestibular gland, scale = 1.0 mm.

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE

OE dis
PRE YAH

FIGURE 3. Chromodoris buchananae sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 8.6 xm. B. Inner lateral
teeth, scale = 27 um. C. Lateral teeth from central portion of half-row, scale = 23.1 um. D. Outer lateral teeth, scale = 25 pm.

narrows abruptly to a curved segment and then enters the short penial bulb, which is adjacent to the
slender vaginal duct at the common gonopore. The distal end of the vas deferens is devoid of any
penial hooks. The female gland mass consists of the large mucous gland and small membrane and al-
bumen glands. Near the exit of the mucous gland there is a small, ovoid vestibular gland. A relatively
short vagina leads to a spherical bursa copulatrix. Adjacent to the vagina, a short duct emerges and

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connects to the pyriform receptaculum seminis. The uterine duct emerges a quarter of the length along
the duct to the receptaculum seminis. The uterine duct 1s relatively short and enters the female gland
mass near the albumen gland.

DISCUSSION. — The color pattern of this species is reminiscent of some members of the
Chromodoris quadricolor complex (Rudman 1982; Gosliner and Behrens 1998). Members of this
group have longitudinal lines, an orange marginal or submarginal band and orange gills and
rhinophores. Of the numerous species in this complex, only C. africana Eliot, 1904 and C. magnifica
(Quoy and Gaimard, 1832) have white lines on the body. The remaining species all have black lines or
other markings. Both C. magnifica and C. africana differ from C. buchananae in having a black rather
than a cream body color. The only other chromodorids with a body color similar to C. buchananae are
C. decora (Pease, 1860) and C. /ekker Gosliner, 1994. Both of these species have a creamy white body
with an orange marginal band and opaque white markings on the notum. In C. decora there is a central
white line that bifurcates anterior to the gills. In C. /ekker there are scattered opaque white spots. Both
species differ from C. buchananae in having opaque white spots and dark plum to black spots
submarginally. They also have white gills and rhinophores whereas C. buchananae has orange gills
and rhinophores and has small brown spots on the notum that are absent in the other two species.

The mantle glands of Chromodoris buchananae are highly ramified as have been described for
members of the C. guadricolor complex (Gosliner and Behrens 1998). The radular teeth of C.
buchananae are similar in configuration to many of the members of this complex, as well. The radular
arrangement differs from that of C. decora and C. lekker in that these species have a well-developed
rachidian row of teeth while in C. buchananae there 1s no trace of a rachidian row. The reproductive
system of C. buchananae does not differ markedly from that of other members of Chromodoris
quadricolor complex. Members of this group of species have a relatively short ejaculatory portion of
the vas deferens and a simple ovoid vestibular gland.

Genus Hypselodoris Stimpson, 1855

Hypselodoris dollfusi (Pruvot-Fol, 1933)
Figs. 1B, 4A—B, SA—D

Glossodoris dollfusi Pruvot-Fol, 1933:126, pl, 1, figs. 7-8; pl. 3, fig. 40.
Hypselodoris dollfusi (Pruvot-Fol, 1933) comb. nov.

MATERIAL EXAMINED. — HOLOTYPE: Muséum National d’Histoire Naturelle, Paris, St. XXV,
Dollfus Expedition, 12 January 1919, leg. R. Ph. Dollfus. CASIZ 127918, one specimen, 21 m depth,
Anemone Gardens, Khor Fakken, Dubai, United Arab Emirates, Gulf of Oman, 24 December 1999,
Carole Harris. CASIZ 127919, one specimen, dissected, 20 m depth, Coral Gardens, Khor Fakken,
Dubai, United Arab Emirates, Gulf of Oman, 16 July 1999, Carole Harris and Leon Betts.

OTHER MATERIAL. — Photographs of additional material from Dubai were examined to deter-
mine variability in external morphology and coloration.

DISTRIBUTION. — Thus far, this species is known only from the type locality, the Gulf of Suez,
and Khor Fakken, United Arab Emirates.

EXTERNAL MORPHOLOGY. — The living animals (Fig. 1B) reach 45-50 mm in length. The body
is opaque white overall, with a bright yellow-orange marginal band. The notum has a series of deep
pink-purple rings of varying diameters, which surround a circle of lavender. In some specimens the
circle within these rings have a bluish tinge. A similarly colored ring is found around the branchial
plume pit and at the base of the rhinophores. The pink rings are situated along the notal margin, with
one at both the anterior and posterior ends and midlaterally between the rhinophores and gill. The
rings vary innumber from 9 to 21. The hyponotum bears 4—6 pink rings and there is one at the tip of the

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE 117

hea

?
oem fae

FiGURE 4. Hypselodoris dollfusi (Pruvot-Fol, 1933) A. Subcutaneous glandular network. B. Reproductive system,
am=ampulla, be=bursa copulatrix, ej=ejaculatory duct, fgm=female gland mass, p=penis, pr = prostate,
Ts = receptaculum seminis, vg = vestibular gland, scale = 1.0 mm.

posterior end of the foot. The rhinophores and gill are red. The rhinophores are perfoliate with 27
lamellae. The branchial plume is partially spreading, with 10 unipinnate gill leaflets.

MANTLE GLANDS. — (Fig. 4A) There are about 22 large posterior glands. A lateral series of
glands is absent. There are 12—18 anterolateral glands on either side of the head. In the paratype, the 18
glands on the right side of the head were smaller (some indicating recent regeneration) than the 13
glands on the left side.

BUCCAL ARMATURE. — The muscular portion of the buccal mass is approximately equal in
length to the oral tube. At the anterior end of the muscular portion of the buccal mass there is a
chitinous labial cuticle, which bears numerous jaw rodlets. The rodlets (Fig. SA) have a short base and
a curved apex without lateral rodlets. The radular formula is 66 x 88.0.88. There is no trace of a
rachidian row of teeth. The inner lateral teeth (Fig. 5B) have a short wide base with a bifid cusp. On the
right side of the radula, inner and outer denticles are absent, while on the left side a single short inner
denticle is present. The remainder of the inner and middle lateral teeth (Fig. 5C) are broad with a bifid
cusp and no lateral denticles on either the inner or outer margins of the teeth. The outer 4—10 teeth (Fig.
5D) also have a bifid cusp and 4—6 denticles on the outer side.

REPRODUCTIVE SYSTEM. — (Fig. 4B) The arrangement of the organs is triaulic. The ampulla is
elongate and curved. It divides into the short, thick, slightly convoluted prostate and short oviduct,

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 5. Hypselodoris dollfusi (Pruvot-Fol, 1933) Scanning electron micrographs. A. Jaw rodlets, scale = 10 um. B. Inner
lateral teeth, scale =38 um. C. Lateral teeth from central portion of half-row, scale=30 um. D. Outer lateral teeth,
scale = 30 pm.

which enters the female gland mass. The prostate narrows into the moderately short ejaculatory por-
tion, which terminates in a slightly enlarged penis. The vaginal duct is narrow and long. The minute,
short, pyriform receptaculum seminis has a short duct which attaches to the vaginal duct at the base of
the spherical bursa copulatrix. The uterine duct is long and narrow forming a loop before entering the

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE 119

female gland mass below the entrance of the oviduct into the mass. It branches from the vagina just be-
low the common insertion of the bursa and receptaculum seminis.The female gland mass is large and
completely developed. A large, lobate vestibular gland is present.

DISCUSSION. — Glossodoris dollfusi was originally described from a single 30 mm preserved
specimen collected from the Gulf of Suez, Egypt. This species has not been documented in other pub-
lications since its original description and its status has remained in question. Pruvot-Fol (1951) sug-
gested that the color description of G. dollfusi was based on a preserved specimen. This statement led
Rudman (1973:196—197) to suggest that the animal was a species of Chromodoris similar to C.
inornata Pease, 1871. Later, Rudman (1983:169) noted the discrepancy between Pruvot-Fol’s origi-
nal description and subsequent comments regarding the coloration being from a living rather than pre-
served specimen. On the basis that the coloration was from a living specimen, Rudman concluded that
the G. dollfusi was most likely a chromodorid, but that its was doubtful that it could be identified with
any degree of certainty. The original description contains two color plates of the living animal and
several figures of the radular teeth. The color plate clearly shows a whitish animal with red to violet
rings on the notum and a yellowish marginal band. The radular teeth are described and depicted as be-
ing bifid, which is characteristic of members of the genus Hypselodoris. There are some minor differ-
ences in the external color pattern between Pruvot-Fol’s original description and the present
specimens. For example the marking on the notum are described by Pruvot-Fol as spots rather than
circles and there are also some minor differences in the coloration of the rhinophores as to the distribu-
tion of reddish pigment. Nevertheless, the description of the radular teeth, with inner edenticulate
teeth and outer denticulate ones closely matches the form of the radular teeth in the present material.
The partially dissected holotype was re-examined. The buccal mass had been removed for prepara-
tion of the radula. No pigment remains. The external anatomy and arrangement of the mantle glands
are entirely consistent with the specimens examined here from the Gulf of Oman. As no other species
of Hypselodoris has a similar color pattern, there is little doubt that the present specimens are
conspecific with Pruvot-Fol’s species. Although, Pruvot-Fol’s animal was from the Gulf of Suez and
the present material is known from the Persian Gulf, there is sufficient geographical overlap in the
biota as to make this a likelihood.

Hypselodoris dollfusi differs markedly from any described species of Hypselodoris (Gosliner
and Johnson, 1999). It is the only member of the genus known to possess circular rings on the notum.
The general whitish body color is similar to most Indo-Pacific members of the genus.

This species appears to be a member of the clade of Hypselodoris species that contains H. fucata
Gosliner and Johnson, 1999; H. kaname Baba, 1994, H. koumacensis Rudman, 1995 and H. paulinae
Gosliner and Johnson, 1999. All members of this clade share three important synapomorphies: erect
rather than spreading branchial plume, a short jaw element shaft and a receptaculum seminis that in-
serts at the base of the bursa copulatrix rather than more distally along the vaginal duct. Hypselodoris
dollfusi shares all of these features with the other members of this clade. In the above-mentioned spe-
cies the mantle glands are small and arranged uniformly around the mantle margin. In H. dollfusi there
are small glands that are interrupted in the mid-region of the body, on either side. The body color of H.
dollfusi most closely resembles that of H. paulinae, but differs in having pink rings rather than red
blotches. Hypselodoris dollfusi also lacks the purple submarginal line on the foot that is present in H.
paulinae.

The inner lateral radular teeth of H. dollfusi closely resemble those of H. fucata and H. paulinae
where a denticle is present on the inner side of the radular tooth, but is absent on the outer side. In H.
kaname and H. koumacensis the outer side of the tooth also bears a denticle. The denticulation of the
radular teeth of H. dol/fusi is most similar to that of H. fucata. In this species only the outer 15 teeth
possess denticles on the outer face of the teeth whereas in H. dollfusi the outer 4—10 teeth are
denticulate. The other members of this clade have more denticulate teeth with the most extreme case
being H. kaname, where all radular teeth are denticulate.

120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 10

The reproductive system of H. dollfusi is similar to that of other members of the clade described
above. All members of this clade have a receptaculum seminis that inserts at the base of the bursa
copulatrix rather than more distally along the vaginal duct. In H. dollfusi, the vagina is narrower than
in the other members of the clade. Also, the ejaculatory portion of the vas deferens appears shorter in
H. dollfusi than in other members of the clade.

Hypselodoris babai sp. nov.
Figs. 1C, 6A-E, 7A—E

TYPE MATERIAL. — HOLOTYPE: CASIZ 115758, one specimen, 55 m depth, Seragaki, Oki-
nawa, 26 April 1996, Bob Bolland. PARATYPES: CASIZ 115804, one specimen, 44 m depth, Seragaki,
Okinawa, 4 April 1997, Bob Bolland. CASIZ 127924, one specimen, 58 m depth, Seragaki, Okinawa,
16 May 2000, Bob Bolland.

ETYMOLOGY. — Hypselodoris babai is named for our friend and colleague Dr. Kikutaro Baba.
He has been an inspiration to us throughout our careers in opisthobranch systematics. For more than
almost 70 years, Baba has been carefully documenting the remarkable diversity of Japanese opistho-
branchs. This 1s a truly remarkable achievement.

DISTRIBUTION. — Thus far, this species is known only from the type locality, Seragaki, Oki-
nawa.

EXTERNAL MORPHOLOGY. — The living animals (Fig. 1C) reach at least 25 mm in length. The
body is opaque white with a rich red-brown central region. There are a series of long irregular oval
white spots distributed over the mantle. The oval marking, beginning between the rhinophores varies
in length and shape, and may extend mid-dorsally half the length of the mantle to the gill. A wide white
band, similar to that on the mantle, edges the foot. A series of white ovals occur medially along the
posterior end of the foot. The gill and rhinophores are bright red. The rhinophores are very long, bear-
ing 25 lamellae. The branchial plume is also very tall and unipinnate, with 5 gill leaflets.

MANTLE GLANDS. — (Fig. 6A, B) The mantle glands are limited to the anterolateral and poste-
rior ends of the dorsum. There are 3-4 large posterior glands and 4—6 smaller, anterolateral glands.
The holotype has three small glands positioned just anterior to the gill plume, on the left side of the
body.

DIGESTIVE SYSTEM AND BUCCAL ARMATURE. — The buccal mass (Fig. 6C) is very small rela-
tive to the size of the body. The oral tube is extremely narrow and elongate. The cerebral nerve ring is
situated well behind the posterior end of the buccal mass. The muscular portion of the buccal mass is
approximately equal in length to the oral tube (Fig. 6D). At its posterior end there are a pair of short
salivary glands and more ventrally situated are the paired buccal ganglia. At the anterior end of the
muscular portion of the buccal mass 1s a chitinous labial cuticle, which bears numerous jaw rodlets.
The rodlets (Fig. 7A) have an elongate base with unifid, bifid and trifid cusps. The radular formula of
the paratype is 57 x 43-45.0.43-45S. There is no trace of a rachidian row of teeth. The innermost lat-
eral teeth (Fig. 7B) have a simple, bifid cusp with one or two denticles on the inner side of the primary
cusp. There are no denticles on the outer side of the cusps. The subsequent inner lateral teeth lack
denticles on either side of the cusp. At approximately the fifteenth row of teeth, a single denticle is
present below the primary cusps. More external midlateral teeth have 4—5 denticles below the cusps
(Fig. 7C). The outermost laterals (Fig. 7D) bear 7-10 rounded denticles below the primary cusps.

REPRODUCTIVE SYSTEM. — (Fig. 6E) The arrangement of the organs is triaulic. The ampulla is
elongate and swollen. It divides into the short, thick, slightly convoluted prostate and short oviduct,
which enters the female gland mass. The prostate narrows into the moderately short ejaculatory por-
tion, which terminates in a much-enlarged penis. The vaginal duct is relatively straight and short. The
small, short, pyriform receptaculum seminis has a short duct that attaches to the vaginal duct near the
middle of its length. The uterine duct is long and narrow, forming several convoluted loops before en-

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE 121

A

dg/ot cns bm

FIGURE 6. Hypselodoris babai sp. nov. A. Subcutaneous glandular network (CASIZ 115758). B. Subcutaneous glandular
network (CASIZ 115804). C. Digestive system, bm = buccal mass; cns = central nervous system; dg/ot = digestive gland/
ovotestis, scale = 5.0 mm. D. Buccal mass, bg = buccal ganglia; mp = muscular portion of buccal mass; ot = oral tube;
ra = radular sac; sg = salivary gland, scale = 1.0 mm. E. Reproductive system, am = ampulla, bc = bursa copulatrix, ej = ejac-

ulatory duct, fgm= female gland mass, p= penis, pr= prostate, rs=receptaculum seminis, vg = vestibular gland,
scale = 1.0 mm.

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Volume 52, No. 10

FIGURE 7. Hypselodoris babai sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 5 um. B. Inner lateral teeth,
scale = 17.6 um. C. Lateral teeth from central portion of half-row, scale = 13.6 um. D. Outer lateral teeth, scale = 12 um.

tering the female gland mass below the entrance of the oviduct into the mass. The uterine duct
branches from the vagina near the base. The female gland mass is large and completely developed. A
large, lobate vestibular gland is present.

DISCUSSION. — This species is most similar in its external coloration to Hypselodoris bullocki
(Collingwood, 1881). Both species have a reddish or pink to purple body color and red to orange gills

GOSLINER AND BEHRENS: NEW SPECIES OF CHROMODORIDIDAE 123

and rhinophores. Hypselodoris babai differs from H. bullocki in having a broader white marginal
band and numerous white spots and blotches over the dorsal surface of the mantle and foot. While de-
tails of the anatomy of H. bullocki remain largely undescribed, the present species differs in several
significant regards. Hypselodoris bullocki has a highly elevated gill sheath and lacks any mantle
glands around the margin of the mantle (present study). Hypselodoris babai has a slightly elevated gill
sheath and has three to four large posterior glands and smaller lateral and anterolateral glands. Details
of the internal anatomy of H. bullocki need to be described to more fully compare these two species.
The present species also bears some resemblance to Durvilledoris pusilla (Bergh, 1874). The latter
species differs externally in having a yellow rather than white marginal band and also has fewer white
spots than in H. babai.

Hypselodoris babai is unusual among described species of Hypselodoris in that the central ner-
vous system and the buccal bulb are much smaller. It is unclear whether this is a also a characteristic of
H. bullocki and related taxa and certainly warrants further study.

ACKNOWLEDGMENTS

This work was supported by a grant from the National Science Foundation Partnerships for En-
hancing Expertise in Taxonomy program (Phylogenetic systematics of dorid nudibranchs) award
9978155, made to the senior author. We thank Robert Bolland, Carol and Neil Buchanan, Carole Har-
ris and Leon Betts for collecting the material studied here. Their enthusiasm has greatly facilitated dis-
covery of many new opisthobranch species in the Indo-Pacific. Virginie Héros of the Muséum
d’Histoire Naturelle, Paris, kindly provided the holotype of Glossodoris dollfusi. We also thank
Angel Valdés for providing valuable comments for improving the manuscript and for preparing the fi-
nal plates and Bill Rudman for providing the critical identification of Hypselodoris dollfusi.

LITERATURE CITED

BABA, K. 1994. Descriptions of four new, rare, or unrecorded species of Hypselodoris (Nudibranchia:
Chromodorididae) from Japan. Venus 53(3):175—187.

1995. Anatomical and taxonomical review of four blue patterned species of Hypselodoris
(Nudibranchia: Chromodorididae) from Japan. Venus 54 (1):1—15.

GOSLINER, T. M. 1994. New species of Chromodoris and Noumea (Nudibranchia: Chromodorididae) from the
western Indian Ocean and southern Africa. Proceedings of the California Academy of Sciences
48(12):239-252.

GOSLINER, T. M. AND D. W. BEHRENS. 1998. Five new species of Chromodoris (Mollusca: Nudibranchia:
Chromodorididae) from the tropical Indo-Pacific Ocean. Proceedings of the California Academy of Sci-
ences 50(5):139-165.

GOSLINER,T. M. AND R. F. JOHNSON. 1999. Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with
a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species.
Zoological Journal of the Linnean Society 125:1—114.

HAMATANI, I. 1995. Two species of Chromodorididae (Nudibranchia), one newly recorded and one newly estab-
lished, from middle Japan. Venus 54(2):101—107.

JOHNSON, R. F. AND T. M. GOSLINER. 1998. The genus Pectenodoris (Nudibranchia: Chromodorididae) from the
Indo-Pacific, with the description of a new species. Proceedings of the California Academy of Sciences
50(12):295—306.

PRUVOT-FOL, A. 1933. Mission Robert Dollfus en Egypte. Opisthobranchiata. Memoires de |’ Institute d’ Egypte
21:89-159.

. 1951. Revision du genre Glossodoris Ehrenberg. Journal de Conchyliologie 91:76—164.

RUDMAN, W. B. 1973. Chromodorid opisthobranch Mollusca from the Indo-West Pacific. Zoological Journal of
the Linnean Society 52(3):175—199.

124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 10

. 1982. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris

quadricolor, C. lineolata and Hypselodoris nigrolineata colour groups. Zoological Journal of the Linnean

Society 76(3):183-241.

. 1983. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris

splendida, C. aspersa and Hypselodoris placida colour groups. Zoological Journal of the Linnean Society

78(2):105-173.

. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: a review of the gen-

era. Zoological Journal of the Linnean Society 81(2 and 3):115—273.

. 1986. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Noumea purpurea

and Chromodoris decora colour groups. Zoological Journal of the Linnean Society. 86:309-353.

. 1987. The Chromodorididae (Opisthobranchia, Mollusca) of the Indo-West Pacific: Chromodoris

epicuria, C. aureopurpurea, C. annulata, C. coi and Risbecia tryoni colour groups. Zoological Journal of

the Linnean Society 90(4):305—407.

. 1995. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: further species
from New Caledonia and the Noumea romeri group. Molluscan Research 16:1—43.

SCHRODL, M 1999. Glossodoris charlottae, a new chromodorid nudibranch from the Red Sea (Gastropoda,
Opisthobranchia). Vita Marina 46(3-4):89—94.

VaLpEs, A., E. MOLLO, AND J. ORTEA, 1999. Two new species of Chromodoris (Mollusca, Nudibranchia,
Chromodorididae) from southern India, with a redescription of Chromodoris trimarginata (Winkworth,
1946). Proceedings of the California Academy of Sciences 51(3):461—-472.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park _
San Francisco, California 94118

~

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 11, pp. 125-142, 12 figs. DECed4 2000 _ December 7, 2000

A Review of the Genus Hemibagrus in Southern Asia, with
Descriptions of Two New Species

by
Heok Hee Ng!

Department of Biological Sciences, National University of Singapore,
10 Kent Ridge Crescent, Singapore 119260

and

Carl J. Ferraris, Jr.
Department of Ichthyology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

The species of the southern Asian bagrid catfish genus Hemibagrus are reviewed in this
study. Five previously-named species are recognized as valid: Hemibagrus maydelli
(Rossel, 1964), H. menoda (Hamilton, 1822), H. microphthalmus (Day, 1877), H. peguensis
(Boulenger, 1894) and H. punctatus (Jerdon, 1849). Two additional species, H. imbrifer and
H. variegatus, from the Salween and Tenasserim River drainages, respectively, are de-
scribed here as new. The status of controversial names Pimelodus menoda Hamilton, 1822,
and Bagrus corsula Valenciennes, 1840, are stabilized with the designation of a single
neotype for both names.

Bleeker (1862) established the genus Hemibagrus for a group of bagrid catfishes characterized
by having a depressed head, rugose head shield not covered by skin, slender occipital process, and
moderately long adipose fin. However, workers since Gunther (1864) have placed species of this ge-
nus in either Mystus Scopoli, 1777, or Macrones Dumeril, 1856, and it was not until Mo’s (1991)
phylogenetic study of the Bagridae that the genus was considered distinct from Mystus.

In much of southern Asia (defined in this study as consisting of the Indian subcontinent and
Myanmar west of the Tenasserim and Salween River drainages) and particularly in India,
Hemibagrus species appear to be less common than in Southeast Asia, inasmuch as they are less fre-
quently encountered in markets, certainly less so than other genera of large bagrid catfishes, such as
Sperata and Rita. In this study, the taxonomy of the southern Asian species of Hemibagrus is re-
viewed and seven valid species, two of which are new and described herein, are recognised.

MATERIALS AND METHODS

Measurements were made point to point with dial calipers and data recorded to tenths of a milli-
meter. Counts and measurements were made on the left side of specimens whenever possible. Sub-
units of the head are presented as percent of head length (HL). Head length and measurements of body
parts are given as percent of standard length (SL). Measurements and counts were made following Ng
and Ng (1995) with the following exceptions: head length is measured from the tip of the snout to the

1Current address: Fish Division, Museum of Zoology, University of Michigan, 1109 Geddes Ave., Ann Arbor, MI 48109

125

126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

posteriormost extremity of fleshy opercular flap. Length of the adipose-fin base is measured from the
anteriormost point of origin to the posteriormost point of the adipose-fin base. Post-adipose distance
is measured from the posteriormost point of the adipose-fin base to the posterior margin of the hypural
complex.

The following additional measurements were made: predorsal, preanal, prepelvic and
prepectoral lengths are those measured from the tip of the snout to the anterior bases of the dorsal,
anal, pelvic, and pectoral fins, respectively. Pelvic- and pectoral-fin lengths are measured from the or-
igin to the tip of the longest ray. Dorsal and pectoral spine lengths are measured from the base to the
tip. Dorsal to adipose distance is measured from the base of the last dorsal-fin ray to the origin of the
adipose fin. Caudal-fin length is the length of the longest ray of the lower lobe measured from the pos-
terior margin of the hypural complex. The length of the caudal peduncle is measured from base of the
last anal-fin ray to the posterior margin of the hypural complex. Nasal-, maxillary-, and mandibu-
lar-barbel lengths are measured from the base to the tip.

Fin-ray counts were obtained under a binocular dissecting microscope using transmitted light.
Vertebral counts were taken from radiographs. Following the method of Roberts (1994), the first ver-
tebra bearing fully-developed ribs was counted as vertebra 6, and the first postanal vertebra is taken to
be the anteriormost vertebra having its hemal spine posterior to the anteriormost anal-fin
pterygiophore. The number in parentheses following a particular count indicates the number of exam-
ined specimens with that count. Drawings of the specimens were made with a Nikon SMZ-10 camera
lucida. Institutional codes for the repositories of specimens follow Eschmeyer (1998).

SYSTEMATIC ACCOUNTS

Hemibagrus imbrifer sp. nov.
Fig. 1

TYPE MATERIAL. — HOLOTYPE: ZRC 45406, 186.6 mm SL; Thailand, Tak Province, Salween
basin, Mae Nam Moei at Ban Wa Le (16°17'24"N, 98°42'21"E); K. Kubota, Apr 1998. PARATYPE:
CMK 13445 (1, 144.2 mm SL), Thailand, Tak Province, Salween basin, Mae Nam Moei at Na Rei
(16°17'23"N, 98°42'20”E); K. Kubota, Mar 1997.

DIAGNOSIS. — Hemibagrus imbrifer can be distinguished from its congeners in having rela-
tively large sensory pores arranged in vertical columns along the sides of the body and the following
unique combination of characters: length of caudal peduncle 18.8—19.5 %SL, interorbital distance
31.7-32.3 %HL, eye diameter 17.3—18.5 %HL, 48 vertebrae (with 24 postanal vertebrae) and 14 gill
rakers on the first gill arch.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. Adipose fin with long base, spanning most of postdorsal distance. Sensory pores
of lateral line system readily visible, arranged in nine vertical columns on sides of body. In %SL: head
length 26.6—28.4, head width 18.8—20.0, head depth 13.0—13.1, predorsal distance 38.8—39.2, preanal
length 68.4~70.6, prepelvic length 51.0-51.5, prepectoral length 23.7—24.5, body depth at anus
14.4-14.9, length of caudal peduncle 18.8—19.5, depth of caudal peduncle 9.8—11.1, pectoral-spine
length 12.2—12.3, pectoral-fin length 16.6, dorsal-spine length 11.7—12.5, length of dorsal fin
22.623.5, length of dorsal-fin base 14.4—16.9, pelvic-fin length 14.6—-15.5, length of anal-fin base
11.1—13.2, caudal-fin length 19.7—21.6, length of adipose-fin base 38.444.2, adipose-fin maximum
height 4.25.5, post-adipose distance 8.1—10.1; in %HL: snout length 39.3—39.8, interorbital distance
31.7-32.3, eye diameter 17.3-18.5, nasal barbel length 41.3-41.7, maxillary barbel length
175.4—211.7, inner mandibular barbel length 48.8—52.0, outer mandibular barbel length 85.4-88.1.

NG AND FERRARIS: HEMIBAGRUS SPECIES 127

FIGURE 1. Hemibagrus imbrifer, ZRC 45406, holotype, 186.6 mm SL; Thailand: Mae Nam Moei.

Branchiostegal rays 10 (1) or 11 (1). Gill rakers 3 + 11 = 14 (1). Vertebrae 26 + 21 = 47 (1) or
26 + 22 = 48 (1).

Fin ray counts: dorsal IT,7 (2); pectoral I,10 (1) or I,10,i(1); pelvic 1,5 (2); anal iv,8 (1) or iv,8,i (1);
caudal i,7,7,i (1) or i,7,8,i (1). Dorsal-fin origin nearer to tip of snout than to caudal flexure. Dorsal
spine stout, without serrations on posterior edge. Pectoral spine stout, with 12—13 large serrations on
posterior edge. Caudal fin forked; distal margins of upper and lower lobes rounded.

COLOR. — Dorsal surface of head and body uniform gray; ventral surfaces of head and body dirty
white; adipose fin gray, distal edge fading to light gray; caudal and anal fins gray, with melanophores
more dense on the fin rays. Distal two-thirds of pectoral and pelvic fins gray, with melanophores more
dense on fin rays and proximal third dirty white.

ETYMOLOGY. — From the Latin imbrifer, meaning rainy. In allusion to the pattern of the sensory
pores being arranged in vertical columns on the sides of the body.

DISTRIBUTION. — Known only from the Salween River drainage (Fig. 2).

REMARKS. — Hemibagrus imbrifer can be differentiated from its congeners in having relatively
large sensory pores of the lateral line system arranged in vertical columns along the sides of the body.
No other species of Hemibagrus have the sensory pores of the lateral line system so obviously visible.
Furthermore, H. imbrifer is one of the only two known species of southern Asian Hemibagrus (the
other being H. variegatus) which has a long-based adipose fin spanning nearly all of the postdorsal
distance. Hemibagrus olyroides from Borneo and all East Asian species allied with H. guttatus have
similar long-based adipose fins, but can be differentiated from H. imbrifer in having more vertebrae
(52-60 vs. 47-48). Hemibagrus baramensis and H. sabanus (both from Borneo) also have long-based
adipose fins with a relatively low vertebral count (44-47), but can be differentiated from H. imbrifer
in lacking the readily-visible sensory pores arranged in vertical columns. Hemibagrus imbrifer can be
differentiated from H. variegatus in having a shorter caudal peduncle (18.8—19.5 %SL vs. 20.2%SL)
with fewer postanal vertebrae (21—22 vs. 24), smaller eyes (17.3—18.5 %HL vs. 23.1%HL), a larger
interorbital distance (31.7—32.3 %HL vs. 28.6%HL), fewer gill rakers (14 vs. 21), and a gray body
with the sensory pores plainly visible (vs. a variegated brown body with the sensory pores not readily
apparent).

128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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0 200 400 600 km
—— ey

65°E 70°

FIGURE 2. Map of southern Asia showing the distribution of Hemibagrus species: H. imbrifer (@); H. maydelli (WM), H.
menoda (W); H. microphthalmus (A); H. peguensis (*); H. punctatus (#*) and H. variegatus (#).

Hemibagrus maydelli (Rossel, 1964)
Fig. 3

Mystus maydelli Rossel, 1964:149, fig. 1; Wilkens 1977:159.

Mystus sp. — Govind and Rajagopal, 1975:79.

Mystus malabaricus (in part) — Jayaram, 1977:32; Talwar and Jhingran, 1991:564.

Mystus krishnensis Ramakrishniah, 1988:139, figs. 1-2; Talwar and Jhingran, 1991:563; Jayaram, 1995:97, 105,
108.

Mystus punctatus (non Jerdon) — Barman, 1993:225, fig. 96.

Hemibagrus maydelli — Grant, 1999:172, fig. 2.

DIAGNOSIS. — Hemibagrus maydelli can be differentiated from its congeners by a unique com-
bination of the following characters: head length 30.8-32.4 %SL, length of caudal peduncle
15.3—16.1 %SL, depth of caudal peduncle 7.8—8.5 %SL, dorsal to adipose distance 4.0—7.0 %SL, eye
diameter 11.5—12.3 %HL, 52 vertebrae, and olive green body with orange fins.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. Head extremely depressed. In %SL: head length 30.8-32.4, head width
18.9-20.5, head depth 11.7—13.3, predorsal distance 42.0-46.7, preanal length 71.1—73.8, prepelvic
length 54.8-58.8, prepectoral length 25.4-28.7, body depth at anus 13.0—15.6, length of caudal

NG AND FERRARIS: HEMIBAGRUS SPECIES 129

FIGURE 3. Hemibagrus maydelli, CAS 62087, 167.2 mm SL; India: Tungabahdra River.

peduncle 15.3—16.1, depth of caudal peduncle 7.8—8.5, pectoral-spine length 13.6—15.3, pectoral-fin
length 17.1—-19.1, dorsal-spine length 11.4—12.9, length of dorsal fin 24.4-27.3, length of dorsal-fin
base 14.4-15.2, pelvic-fin length 13.9-15.5, length of anal-fin base 10.7—13.1, caudal-fin length
19.7-23.2, length of adipose-fin base 14.3—20.3, adipose-fin maximum height 4.0—5.1, post-adipose
distance 14. 1—14.6, dorsal to adipose distance 4.0—7.0; in %HL: snout length 31.1—35.2, interorbital
distance 28.6—29.9, eye diameter 1 1.5—12.3, nasal barbel length 31.9-35.4, maxillary barbel length
237.6—298.9, inner mandibular barbel length 44.1—48.9, outer mandibular barbel length 78.2—-93.9.
Branchiostegal rays 9 (6). Gill rakers 3 + 9 = 12 (1). Vertebrae 27 + 25 = 52 (2).

Fin ray counts: dorsal II,7 (6); pectoral I,7 (1), 1,8 (1) or I,9 (4); pelvic 1,5 (6); anal iv,8 (3), v,8 (2)
or iv,10 (1); caudal 1,7,8,i (6). Dorsal-fin origin nearer to tip of snout than to caudal flexure. Dorsal
spine stout, without distinct serrations on posterior edge. Pectoral spine stout, with 13-19 serrations
on posterior edge. Caudal fin forked; first principal ray of upper lobe extending into a long filament;
distal margin of upper and lower lobes rounded.

COLOR. — Preserved specimens have dorsal surface of head and body brown, gradually fading to
dirty white on ventral surface. Pectoral, pelvic, anal and caudal fins brown with melanophores con-
centrated in interradial membranes. Dorsal fin brown with melanophores evenly distributed. Live
specimens are olive green in color with orange-tipped fins (after Ramakrishniah 1988).

DISTRIBUTION. — Known only from the middle reaches of the Krishna River drainage in south-
ern India (Fig. 2).

REMARKS. — Hemibagrus maydelli can be differentiated from all other species of Hemibagrus
on the Indian subcontinent in having more vertebrae (52 vs. 44-46). In its general morphology, H.
maydelli resembles both H. microphthalmus and H. wyckioides in having a strongly depressed head. It
can be differentiated from both species in having a shorter distance between the dorsal and adipose
fins (4.0-7.0 %SL vs. 8.6—-14.2%SL), larger eyes (eye diameter 1 1.5—12.3 %HL vs. 8.4-11.6%HL),
and an olive green body with orange fins (vs. gray body with red fins). Hemibagrus maydelli can be
further differentiated from H. microphthalmus in having a shorter and deeper caudal peduncle (length
of caudal peduncle 15.3-16.1 %SL vs. 16.4-18.1 %SL, depth of caudal peduncle 7.8—-8.5 %SL vs.
6.8-7.7 %SL), a longer head (30.8-32.4 %SL vs. vs. 29.4-31.0 %SL).

Govind and Rajagopal (1975) reported the occurrence of H. maydelli from the Tungabahdra
River as an unidentified the species of Mystus, stating that it resembled H. punctatus and further stud-
ies were needed to clarify its identity. Barman (1993) then erroneously considered H. punctatus to be
present in the Krishna River drainage (H. punctatus is only known from the Cauvery River drainage
further south), basing his record on that of Govind and Rajagopal (1975). Hemibagrus maydelli is a
relatively large species that grows up to 1650 mm TL and 58.5 kg in weight (Govind and Rajagopal
1975; Jayaram 1995).

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MATERIAL EXAMINED.— ZMH 2180 (1), holotype, 82.0 mm SL; India: Maharashtra state,
Bhima River at Wadgaon; Maydell, 1955 (photograph and radiograph examined). ZSI FF2532,
271.8 mm SL; India: Andhra Pradesh State, Krishna River below Nagarjunasagar Reservoir; M.
Ramakrishniah, 10 Feb 1982 (holotype of Mystus krishnensis). ZSI FF 2533 (1,402.6 mm SL), local-
ity as for ZSI FF2532; M. Ramakrishniah, 9 Mar 1983 (paratype of Mystus krishnensis). ZS] FF 2534
(1, 228.0 mm SL), locality as for ZS] FF2532; M. Ramakrishniah, 29 Jan 1985 (paratype of Mystus
krishnensis). ZS] FF 2535 (1, 278.8 mm SL), locality as for ZS] FF2532; M. Ramakrishniah, 14 Dec
1980 (paratype of Mystus krishnensis). CAS 62087 (2, 167.2-214.7 mm SL), India, Karnataka State,
Bellary District, Krishna River basin, Tungabahdra River and reservoir at Hospet, Hampi and
Kampli; T. R. Roberts, 28 Jan—3 Feb 1985.

Hemibagrus menoda (Hamilton, 1822)
Figs. 4, 5

Pimelodus menoda Hamilton, 1822:203, pl. | fig. 72 (figure erroneously labelled Mugil corsula; see below for
explanation).

Bagrus trachacanthus Valenciennes, in Cuvier and Valenciennes, 1840:419; Bleeker, 1853:56.

Bagrus corsula Valenciennes, in Cuvier and Valenciennes, 1840:408; Bleeker, 1853:56.

Macrones menoda — Ginther, 1864:74; Day, 1871b:706 (in part).

Macrones trachacanthus — Ginther, 1864:75.

Macrones corsula — Day, 1869:307; 1877:446, pl. 100 fig. 5; 1889:153 (in part).

Mystus menoda—Shaw and Shebbeare, 1937:92, fig. 91; Jayaram and Singh, 1977:263; Menon, 1977:61; Ataur
Rahman, 1974:7, 1989:199, fig. 119D; Shrestha. 1994:52, fig. 80.

Mystus (Mystus) menoda (in part) — Jayaram, 1954:546, fig. 9.

Mystus (Mystus) menoda trachacanthus — Jayaram, 1954:546.

Mystus (Mystus) punctatus (in part) — Jayaram, 1954:547.

Mystus (Mystus) menoda — Motwani et al., 1962:21; Srivastava, 1968:73, fig. 46.

Mystus corsula — Qureshi, 1965:42, fig. 103.

Mystus menoda menoda—Jayaram, 1977:33, fig. 25B (in part); Sen, 1985:137, fig. 75; 1992:183, fig. 60; Dutta et
al., 1993:26.

Mystus menoda trachacanthus — Jayaram, 1977:33; Singh and Yazdani, 1993:21.

Mystus trachacanthus — Mo, 1991:130.

Hemibagrus menoda — Mo, 1991:132.

DIAGNOSIS. — Hemibagrus menoda can be differentiated from its congeners by the following
unique combination of characters: head length 32.7-33.5 %SL, head depth 14.2—-15.3 %SL, depth of
caudal peduncle 7.5—8.8 %SL, eye diameter 11.9—12.3 %HL, a pattern of dark dots arranged in verti-
cal columns on the sides of the body, a convex snout and a broad, shallowly incised humeral process.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. In %SL: head length 32.7-33.5, head width 19.3-21.7, head depth 14.2—15.3,
predorsal distance 42.2-45.3, preanal length 71.2—77.5, prepelvic length 55.0-60.5, prepectoral
length 28.6-30.3, body depth at anus 13.3—17.1, length of caudal peduncle 15.2—17.0, depth of caudal
peduncle 7.5—8.8, pectoral-spine length 14.6—19.9, pectoral-fin length 18.6—24.0, dorsal-spine length
13.7-16.6, length of dorsal fin 24.3-27.4, length of dorsal-fin base 14.2—16.5, pelvic-fin length
14.7-16.1, length of anal-fin base 11.9—12.6, caudal-fin length 22.8—24.8, length of adipose-fin base
13.0-15.8, adipose-fin maximum height 3.8—4.5, post-adipose distance 15.0—-17.4, dorsal to adipose
distance 14.2—14.9; in %HL: snout length 36.2—38.8, interorbital distance 31.4—35.1, eye diameter
11.9-12.3, nasal barbel length 26.4-37.8, maxillary barbel length 191.4—213.3, inner mandibular bar-

NG AND FERRARIS: HEMIBAGRUS SPECIES 131

FIGURE 4. Hemibagrus menoda, illustration from Hamilton (1822), pl. 1, fig. 72.

bel length 36.8—48.1, outer mandibular barbel length 65.2—73.5. Branchiostegal rays 10 (2) or 11 (1).
Gill rakers 3 + 9 = 12 (1) or 4+ 14= 18 (1). Vertebrae 22 + 22 = 44 (1) or 24 + 21 = 45 (1).

Fin ray counts: dorsal II,7 (4); pectoral I,7 (2) or I,8 (2); pelvic 1,5 (4); anal iv,8 (2), 111,9 (1) or iv,9
(1); caudal 1,7,8,1 (4). Dorsal-fin origin nearer to tip of snout than to caudal flexure. Dorsal spine stout,
with 6—9 serrations on posterior edge. Pectoral spine stout, with | 1—17 serrations on posterior edge.
Caudal fin forked; distal margins of upper and lower lobes rounded.

COLOR. — Preserved specimens have dorsal surface of head and body grayish-brown, gradually
fading to dirty white on ventral surface. Lateral surface of body with about nine vertical columns of

FIGURE 5. Hemibagrus menoda, neotype, UMMZ 208726, 202.6 mm SL; Bangladesh: Shari River.

132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

black spots, largest spots being those in middle of columns along lateral line. Dorsal, pectoral, pelvic,
and anal fins grayish brown, with scattered melanophores on fin rays and interradial membranes. Cau-
dal fin grayish brown, with lighter hue along posterior margin, and on procurrent and outer principal
caudal rays.

DISTRIBUTION. — Known from the Ganges, Brahmaputra, Mahanadi and Godavari river
drainages in Bangladesh and northern India (Fig. 2).

REMARKS. — As indicated by the extensive synonymy of this species, the identity of H. menoda
has been problematic. This is due, in large part, to the brief and vague original description in Hamilton
(1822) and confusion over the significance of an illustration in that publication. The illustration in
question, Plate 1 (Fig. 72), includes a lateral view of a catfish identified as “Mugil corsula.” A
tipped-in corrigenda in one copy of Hamilton (1822) at the California Academy of Sciences includes
the following statement: “For Mugil Corsula read Pimelodus, the Mugil Corsula being delineated
Plate IX, Fig. 97.” This sentence has been interpreted by several authors (e.g., Valenciennes, in
Cuvier and Valenciennes, 1840 and Day, 1877) to mean that the name for the fish should be Pimelodus
corsula, a name that is otherwise not mentioned in Hamilton’s book. By examining the original draw-
ings from which the plates of Hamilton (1822) were made, Day (1871a) found the names Pimelodus
menoda and P. telgagra associated with the figure labeled as drawing no. 18, which was later pub-
lished as Plate 1 (Fig. 72). On this basis, he later (Day 1877) placed the name Pimelodus menoda Ham-
ilton in the synonymy of Pimelodus corsula. Somewhat later, Hora (1929) examined a duplicate set of
drawings prepared for Hamilton and noted that the drawing that formed the basis of Plate 1 (Fig. 72)
was identified as Pimelodes telagra menoda. On the basis of that drawing, Hora also concluded that
the fish illustrated in Plate 1 (Fig. 72) was Pimelodus menoda Hamilton. Following Hora, most Indian
ichthyologists have used the name Macrones menoda, or Mystus menoda, for the species that is repre-
sented in Hamilton’s Plate 1 (Fig. 72). However, Valenciennes (in Cuvier and Valenciennes 1840)
had previously attempted to match that illustration with one of the species described in Hamilton’s
text. Valenciennes concluded that the description of only one species, Pimelodus carcio, resembled
the illustration to any degree. Even so, the description was considered sufficiently different such that
Valenciennes chose not to associate the name with the figure and, instead, adopted the name Bagrus
corsula for the illustrated species.

It is generally recognized that Hamilton did not retain specimens. Therefore, the identity of Ham-
ilton’s Pimelodus menoda and the relationship between that name and Plate | (Fig. 72) remains open
to question. In order to stabilize the name Hemibagrus menoda (Hamilton), we believe it necessary to
name a neotype for Pimelodus menoda and, in keeping with the current use of the name, we designate
UMMZ 208726 as neotype. To further stabilize the nomenclature of this group, we choose the same
specimen as the neotype of Bagrus corsula Valenciennes, a species name based only on Hamilton’s
Plate | (Fig. 72). By this action, Bagrus corsula becomes an objective junior synonym of Pimelodus
menoda.

Hemibagrus menoda differs from all other species of Hemibagrus except H. peguensis in having
a pattern of dark dots arranged in vertical columns on the sides of the body. Hemibagrus menoda dif-
fers from H. peguensis in having a longer head (32.7—33.5 %SL vs. 29.0-32.5 %SL), a more convex
snout (Fig. 6) and a broader, less deeply incised humeral process (Fig. 7). Hemibagrus menoda is
found only in the river drainages in Bangladesh and northern India. All records of H. menoda from
Myanmar refer to H. peguensis instead.

MATERIAL EXAMINED. — NEOTYPE: UMMZ 208726, 202.6mm SL; Bangladesh: Surma
(Meghna) drainage, Sharighat bazaar, 22 miles NE of Sylhet on Sylhet-Shillong highway (said to be
from Shari River); W. J. Rainboth and A. Rahman, 20 Feb 1978. Other material: ANSP 85796 (1,

(13.0 mm SL), India, Bombay; Bombay Natural History Society, 1923. MNHN 1191 (1, 285.4 mm
SL syntype of Bagrus trachacanthus), India, Bengal, A. Duvaucel, date unknown. ZSI 426 (1,
167.2 mm SL), India, Bombay: F. Day collection.

NG AND FERRARIS: HEMIBAGRUS SPECIES 133

a

FIGURE 6. Dorsal views of heads of: a. Hemibagrus peguensis, CAS 133789, 212.7 mm SL; b. H. menoda, UMMZ 208726,
202.6 mm SL.

Hemibagrus microphthalmus (Day, 1877)
Fig. 8

Macrones microphthalmus Day, 1877:446, pl. 100 fig. 4; 1889:154; Vinciguerra, 1890:225.
Mystus (Mystus) menoda microphthalmus — Jayaram, 1954:547.

Mystus microphthalmus —Tint Hlaing, 1971:513; Jayaram, 1977:34; Viswanath and Singh, 1986:197, fig. 1; Mo,
1991:130; Talwar and Jhingran, 1991:566.

Hemibagrus micropthalmus — Ukkatawewat and Vidthayanon, 1998:46.

DIAGNOSIS. — Hemibagrus microphthalmus can be differentiated from its congeners by a
unique combination of the following characters: length of dorsal-fin base 13.7—16.7 %SL, dorsal to
adipose distance 8.6—14.2 %SL, interorbital distance 28.4-31.8 %HL, eye diameter 9.2—11.3 %HL
and a rounded snout.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. Head extremely depressed. In %SL: head length 29.4-31.0, head width
18.0-19.7, head depth 11.2—14.0, predorsal distance 40.6—44.1, preanal length 70.8—74.8, prepelvic
length 53.2-55.9, prepectoral length 25.5—28.9, body depth at anus 10.2—14.5, length of caudal
peduncle 16.4—18.1, depth of caudal peduncle 6.8—7.7, pectoral-spine length 11.0—14.1, pectoral-fin
length 13.9-17.6, dorsal-spine length 9.2—10.8, length of dorsal fin 23.0-25.5, length of dorsal-fin
base 13.7-16.7, pelvic-fin length 13.1—-15.4, length of anal-fin base 11.2—-13.3, caudal-fin length
19.8-23.5, length of adipose-fin base 18.4—25.9, adipose-fin maximum height 4.2—5.8, post-adipose
distance 13.4—16.1, dorsal to adipose distance 8.6—14.2; in %HL: snout length 32.9-34.7, interorbital

134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

a distance 28.4-31.8, eye diameter 9.2-11.3, na-
sal barbel length 24.5—35.3, maxillary barbel
length 256.4-300.6, inner mandibular barbel
length 36.0-57.6, outer mandibular barbel
length 65.8—97.2. Branchiostegal rays 9 (3) or
10 (8). Gill rakers 2+9=11 (1), 2+10=12
(2),3+7=10 (1) or 3+9=12 (1). Vertebrae
26 + 25 = 51 (1), 28 + 23 = 51 (1), 28 + 24 = 52
(2), 28+25=53 (), 29 +25 —540h) eon
30 + 24 =54 (1).

Fin ray counts: dorsal II,7 (11); pectoral 1,7
(1), 1,7,1 C), 18 C1), 18,1 C), Lo @ joni,
pelvic 1,5 (11); anal iv,8 (6), 111,9 (1) or iv,9 (4);

b caudal 1,7,8,i (11). Dorsal-fin origin nearer to tip
of snout than to caudal flexure. Dorsal spine
stout, without distinct serrations on posterior
edge. Pectoral spine stout, with 9-10 serrations
on posterior edge. Caudal fin forked; distal mar-
gins of upper and lower lobes rounded.

COLOR. — Preserved specimens with dor-
sal surface of head and body uniform gray (live
or freshly-dead specimens generally darker, fad-
ing on preservation); ventral surfaces of head

FIGURE 7. Humeral processes of: a. Hemibagrus menoda, and body dirty white; adipose fin gray, distal
UMMZ 208726, 202.6 mm SL; b. H. peguensis, CAS 89005, edge orange in life but fading to light gray on
261.9 mm SL. preservation; caudal fin red in life, fading to gray

with very light gray procurrent and outer princi-

pal caudal-rays on preservation; all other fins gray with distal portions of fin rays and inter-radial
membranes red in life, fading to light gray on preservation. Maxillary barbel white.

DISTRIBUTION. — Known from the Salween River of Thailand, Irrawaddy and Sittang drainages
in Myanmar and the Manipur drainage in India (Fig. 2).

REMARKS. — Hemibagrus microphthalmus is similar in form and coloration to H. wyckioides.
In recent years, Roberts (1993) and Roberts and Warren (1994) have considered the latter species a ju-
nior synonym of H. microphthalmus. However, as discussed in Ng and Rainboth (1999), the two spe-
cies differ in the shape of their snouts: H. microphthalmus has a rounded snout while H. wyckioides
has a truncate snout (Fig. 11). Hemibagrus microphthalmus also has a narrower head (18.0—-19.7 %SL
vs. 19.5-23.9 %SL), shorter dorsal-fin base (13.7—16.7 %SL vs.16.3—18.3 %SL) and more closely-set
eyes (interorbital distance 28.4-31.8 %HL vs. 31.6-36.9 %HL) compared to H. wyckioides. Finally,
the two species are geographically separate: H. microphthalmus is found only in the Salween,
Irrawaddy, Sittang and Manipur drainages in Myanmar and India while H. wyckioides is only known
from the Mekong and Chao Phraya drainages, and possibly the Mae Khlong drainage [reported by
Roberts (1993) as H. microphthalmus, but we have not examined any specimen from the Mae Khlong
to ascertain the exact identity of Roberts’ record] in central Indochina.

MATERIAL EXAMINED. — AMS B.7918 (1, 164.0 mm SL syntype), and ZSI 2952 (1, 138.9 mm
SL syntype), Burma: Irrawaddy River; F. Day, date unknown. BMNH 1893.2.16.7 (1, 133.5 mm SL).
CAS 93192 (3, 132.0-151.5 mm SL), Myanmar: Irrawaddy River drainage, Mandalay markets; T. R.
Roberts, Apr 1993. CMK 14706 (1, 204.6 mm SL), Thailand: Tak province, Mae Nam Moei at Mae
Sarid (17°26'25”, 98°3'41”E); M. Kottelat and K. Kubota, 8 Apr 1998. NRM 13892 (1, 116.1 mm
SL), Myanmar: Mandalay Division, Irrawaddy River drainage, Mandalay area; O. Hetzel, Apr 1935.

NG AND FERRARIS: HEM/IBAGRUS SPECIES 135

FIGURE 8. Hemibagrus microphthalmus, USNM 344670, 201.6 mm SL; Myanmar, Mandalay.

NRM 24979 (2, 144.6-165.1 mm SL), Myanmar: Sagaing Division, Irrawaddy River drainage,
Shweli River; Maung Lu Daw, Feb 1935. NRM 31072 (1, 147.1 mm SL), Myanmar: Yangon Divi-
sion, Yangon River at Yangon; R. Malaise, 30 Nov 1934. USNM 44754 (1, 158.7 mm SL),
Myanmar: Irrawaddy River drainage, Mandalay; L. Fea, 1885-1889. USNM 344670 (2,
201.6—239.8 mm SL), Myanmar: Irrawaddy River drainage, Mandalay fish markets; C. J. Ferraris, D.
Catania and U Myint Pe, 23 Apr 1996.

FiIGuRE 9. Dorsal views of heads of: a. Hemibagrus microphthalmus, NRM 13892, 116.1 mm SL; b. H. wyckioides, UMMZ
213974, 177.9 mm SL.

136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

Hemibagrus peguensis (Boulenger, 1894)
Fig. 10

Bagrus menoda (non Hamilton, 1822)— Blyth, 1860:285.

Macrones menoda (non Hamilton, 1822) — Day, 1871b:706 (in part); Vinciguerra, 1890:223.
Macrones corsula (non Hamilton, 1822) — Anderson, 1879:863; Kyaw Win, 1971:53, fig. 21.
Macrones peguensis Boulenger, 1894:196.

Mystus (Mystus) menoda (in part) — Jayaram, 1954:546.

Mystus (Mystus) peguensis — Jayaram, 1954:552.

Mystus menoda menoda (in part) — Jayaram, 1977:33.

Mystus peguensis — Jayaram, 1977:35; Talwar and Jhingran, 1991:569.

Hemibagrus peguensis — Mo, 1991:132.

DIAGNOSIS. — Hemibagrus peguensis can be differentiated from its congeners by a unique com-
bination of the following characters: head length 29.0-32.5 %SL, a gently curving snout and a slen-
der, deeply-incised humeral process, and nine vertical columns of black spots on the sides of the body.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. In %SL: head length 29.0—32.5, head width 18.8—21.7, head depth 12.9-15.2,
predorsal distance 39.8-44.9, preanal length 70.6—74.0, prepelvic length 52.5—-57.6, prepectoral
length 25.4-30.8, body depth at anus 12.3—16.3, length of caudal peduncle 16.2—18.2, depth of caudal
peduncle 7.6—8.8, pectoral-spine length 15.3—19.6, pectoral-fin length 18.2—20.9, dorsal-spine length
13.5—16.2, length of dorsal fin 23.8-27.8, length of dorsal-fin base 13.6—15.9, pelvic-fin length
13.3—15.5, length of anal-fin base 11.2—14.0, caudal-fin length 19.3—23.9, length of adipose-fin base
14.2—19.3, adipose-fin maximum height 3.74.9, post-adipose distance 15.4—17.4, dorsal to adipose
distance 10.4-15.0; in %HL: snout length 36.3—39.9, interorbital distance 30.5—35.4, eye diameter
11.2—13.5, nasal barbel length 22.7-34.0, maxillary barbel length 160.6—2 12.6, inner mandibular bar-
bel length 34.5—45.1, outer mandibular barbel length 57.4—70.2. Branchiostegal rays 9 (4) or 10 (13).
Gill rakers 3+9=12 (2) or 4+8=12 (1). Vertebrae 23+21=44 (2), 24+20=44 (1),
23 + 22 = 45 (2) or 24 + 21 = 45 (4).

Fin ray counts: dorsal IT,6 (1) or II,7 (16); pectoral I,8 (5), 1,8,i (3), 1,9 (5), 1,9,i (3) or I,10 (1); pel-
vic 1,5 (17); anal iv,6 (1), iv,7 (1), i11,8 (2), iv,8 (7), i11,9 (1), v,8 (2) or iv,9 (3); caudal i,7,8,i (17). Dor-
sal-fin origin nearer to tip of snout than to caudal flexure. Dorsal spine stout, with 6-11 serrations on
posterior edge. Pectoral spine stout, with 15—19 serrations on posterior edge. Caudal fin forked; distal
margins of upper and lower lobes rounded.

COLOR. — Preserved specimens with dorsal surface of head and body grayish brown, gradually
fading to dirty white on ventral surface. Lateral surfaces of body with about nine vertical columns of
black spots, largest spots being those in middle of columns, along the lateral line. Dorsal, pectoral, pel-
vic and anal fins grayish brown, with scattered melanophores on fin rays and interradial membranes.
Caudal fin grayish brown, with lighter hue along posterior margin, and on procurrent and outer princi-
pal caudal rays.

DISTRIBUTION. — Known from the Irrawaddy, Sittang and Pegu drainages in Myanmar (Fig. 2).

REMARKS. — Hemibagrus peguensis has long been misidentified as H. menoda (e.g., Day 1889;
Jayaram 1954). Even in cases where it was considered a distinct species (e.g., Jayaram 1977; Talwar
and Jhingran 1991), no clear distinguishing characters were used to separate the two species, nor was
it recognised that H. peguensis superficially resembled H. menoda. As a result, these accounts often
listed the presence of H. menoda in Myanmar when in fact the records actually refer to H. peguensis.
Hemibagrus peguensis can be differentiated from H. menoda in having a shorter head (29.0—32.5

NG AND FERRARIS: HEMIBAGRUS SPECIES 137

FiGuRE 10. Hemibagrus peguensis, ZRC 43511, 243 mm SL; Myanmar, Yangon Division, Win Paw Hta River.

%SL vs. 32.7-33.5 %SL), a gently curving snout (Fig. 6) and a thinner, more deeply incised humeral
process (Fig. 7).

Hemibagrus peguensis can be differentiated from H. punctatus in having a longer adipose-fin
base (14.2—19.3 %SL vs. 10.1—-13.2 %SL), a shorter distance between the dorsal and adipose fins
(10.4-15.0 %SL vs. 16.3—19.4 %SL), a narrower caudal peduncle (7.6—8.8 %SL vs. 8.8-9.9 %SL)
and a smaller eye (11.2—13.5 %HL vs. 13.8-15.7 %HL).

The original description of H. peguensis gives the total lengths of the syntypes as 20 mm. This is
clearly a typographical error for 20 cm, which is the approximate total length of each of the syntypes.

MATERIAL EXAMINED. —BMNH_ 1894.5.21:25-26 (2, 168.8-185.1 mm SL_ syntypes),
Myanmar, Taungoo; E. W. Oates, 1893. BMNH 1891.11.30:200—209 (16, 168.7-285.6 mm SL),
Myanmar, Sittang River; E. W. Oates, 8 May 1891. CAS 89005 (1, 261.9 mm SL), Myanmar: Bago
Division, Sittang River at Taungoo; C. J. Ferraris and D. Catania, 7 Apr 1996. CAS 93201 (1,
148.0 mm SL), Myanmar, Irrawaddy River drainage, Mandalay markets; T. R. Roberts, Apr 1993.
CAS 133789 (1, 212.7 mm SL), Myanmar, Yangon Division, Pegu River drainage, 9 miles NW of
Hlegu; A. W. Herre, 2 Apr 1937. NRM 15064 (2, 116.8—138.9 mm SL), Myanmar, Sagaing Division,
Irrawaddy River drainage, Shweli River; Maung Lu Daw, Feb 1935. NRM 15105 (1, 166.8 mm SL),
Myanmar, Mandalay Division, Mandalay; collector unknown, 1935. NRM 31068 (1, 186.1 mm SL),
Myanmar, Kachin State, Irrawaddy River drainage, Myitkyina; R. Malaise, 10 Mar 1934. NRM
39397 (1, 290.6 mm SL), Myanmar, Bago Division, Bago; R. Malaise, 1934. ZSI 550 (1, 241.2 mm
SL) and ZSI 551 (1, 265.6 mm SL), Myanmar: Tagoung; J. Anderson, date unknown.

Hemibagrus punctatus (Jerdon, 1849)
Fig. 11

Bagrus punctatus Jerdon, 1849:339.

Hemibagrus punctatus — Day, 1867:284.

Macrones punctatus — Day, 1877:445, pl. 100 fig. 3; 1889:153.

Mystus (Mystus) punctatus (in part) — Jayaram, 1954:547.

Mystus punctatus — Jayaram, 1977:36, fig. 25A; 1981:197, 201, fig. 95A; Mo, 1991:131; Talwar and Jhingran,
1991:570, fig. 188.

Mystus menoda menoda (non Day) — (?)Barman, 1993:223, fig. 94.

DIAGNOSIS. — Hemibagrus punctatus can be differentiated from its congeners by a unique com-
bination of the following characters: head length 28.1—29.6 %SL, head depth 11.9-14.3 %SL, depth
of caudal peduncle 8.8—9.9 %SL, eye diameter 13.8—15.7 %HL.

138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. In %SL: head length 28.1—29.6, head width 16.4—20.5, head depth 11.9-14.3,
predorsal distance 39.8—42.0, preanal length 71.0—74.2, prepelvic length 53.3—54.2, prepectoral
length 26.2—26.6, body depth at anus 1 1.9—14.3, length of caudal peduncle 16.1—18.6, depth of caudal
peduncle 8.8—9.9, pectoral-spine length 15.6—18.1, pectoral-fin length 18.7—21.1, dorsal-spine length
13.9-15.9, length of dorsal fin 24.7—27.6, length of dorsal-fin base 14.7—17.4, pelvic-fin length
14.9-17.2, length of anal-fin base 11.8—14.3, caudal-fin length 21.4—23.9, length of adipose-fin base
10.1—-13.2, adipose-fin maximum height 3.9—5.4, post-adipose distance 16.2, dorsal to adipose dis-
tance 16.3—19.4; in %HL: snout length 35.7—38.9, interorbital distance 31.3—32.5, eye diameter
13.8—-15.7, nasal barbel length 27.5—40.3, maxillary barbel length 163.2—203.4, inner mandibular bar-
bel length 31.7—45.6, outer mandibular barbel length 68.8—80.2. Branchiostegal rays 9 (2) or 10 (1).
Gill rakers 4 + 8 = 12 (1) or 5 + 13 = 18 (1). Vertebrae 25 + 21 = 46 (1).

Fin ray counts: dorsal II,7 (3); pectoral I,9 (2) or I,10 (1); pelvic 1,5 (3); anal iv,8 (1) or iv,9 (2);
caudal i,7,8,i (3). Dorsal-fin origin nearer to tip of snout than to caudal flexure. Dorsal spine stout,
with 5—7 serrations on posterior edge. Pectoral spine stout, with 12—19 serrations on posterior edge.
Caudal fin forked; distal margins of upper and lower lobes rounded.

COLOR. — Preserved specimens have dorsal surface of head and body grayish brown, fading to
dirty white on ventral surface. Lateral surface of body with about 9—10 black spots arranged in hori-
zontal row along lateral line. Dorsal, pectoral, pelvic and anal fins grayish brown, with scattered
melanophores on fin rays and interradial membranes. Caudal fin grayish brown, with lighter hue
along posterior margin, and on procurrent and outer principal caudal rays.

DISTRIBUTION. — Known only from the Cauvery River drainage in southern India (Fig. 2).

REMARKS. — Hemibagrus punctatus has generally been regarded as a species of Mystus, as re-
cently as the work of Mo (1991). Our examination of specimens shows that the species has the de-
pressed head characteristic of Hemibagrus, and should be placed within this genus instead.

Hemibagrus punctatus differs from H. menoda in having a shorter, flatter head (head length
28.1—29.6 %SL vs. 32.7-33.5%SL; head depth 11.9-14.3 %SL vs. 14.2-15.3%SL), deeper caudal
peduncle (8.8—9.9 %SL vs. 7.5—8.8%SL) and larger eye (13.8-15.7 %HL vs. 11.9-12.3%HL). The
color pattern of H. punctatus differs from that of H. menoda and H. peguensis. In the latter two spe-
cies, the sides of the body are marked with a series of vertical columns of black spots, the largest of
which is in the middle of the columns along the lateral line whereas in H. punctatus, there is only a sin-
gle row of black spots located along the lateral line.

Babu Rao and Chattopadhyay (1969) record H. punctatus from west Bengal based on a specimen
of 62.0 mm SL. According to their description, the specimen lacked the black spots on the sides of the
body, a feature they attributed to the small size of the specimen. We have not examined enough mate-
rial to ascertain if this is indeed the case, but the rest of their description does not seem to match that of
H. punctatus. They stated that the maxillary barbels reached up to the middle of the pelvic fins, but the
specimens of H. punctatus we examined do not have the maxillary barbels extending beyond the ori-
gin of the pelvic fins. Furthermore, they describe the snout of their specimen as being narrow (com-
pared to Mystus gulio), but the snout of H. punctatus is actually broader than that of M. gulio.
Therefore in the light of the available evidence, it seems very unlikely that their specimen was really
H. punctatus. We have also examined specimens recorded as H. punctatus from Bombay, and have
reidentified them as H. menoda. Barman (1993) recorded H. menoda (as Mystus menoda menoda)
from the Krishna River drainage; although he had not examined any specimens, we feel that his record
may refer to H. punctatus instead, given the proximity of the Cauvery and Krishna river drainages.
Therefore, on the basis of the specimens we have examined and the literature, it appears that the distri-

NG AND FERRARIS: HEMIBAGRUS SPECIES 139

FIGURE 11. Hemibagrus punctatus, ZS] FF 1223, | ex., 193.1 mm SL; India: Hemavathy River at Huliva Laom.

bution of H. punctatus is restricted to the Cauvery River drainage (although it may occur in the
Krishna River drainage) in southern India.

MATERIAL EXAMINED. — BMNH 1868.5.14:8 (1, 154.6 mm SL), India; F. Day collection. - ZSI
F12403 (1, 120.2 mm SL), India, Karnataka State, Cauvery River at Coorg; C. R. Narayan Rao. ZSI
FF 1223 (1, 193.1 mm SL), India, Karnataka State, Hemavathy River at Huliva Laom; K. C. Jayaram,
7 May 1977.

Hemibagrus variegatus sp. nov.
Fig.12

TYPE MATERIAL. — HOLOTYPE: BMNH 1992.11.16:11, 121.2 mm SL; Myanmar: Tenasserim
River; T. R. Roberts, 3—8 Mar 1992.

DIAGNOSIS. — Hemibagrus variegatus can be differentiated from its congeners by a unique
combination of the following characters: length of caudal peduncle 20.2 %SL, length of adipose-fin
base 30.8 %SL, eye diameter 23.1 %HL, interorbital distance 28.6 %HL, 21 gill rakers, 50 vertebrae
(24 postanal) and a variegated brown body with the sensory pores not readily visible.

DESCRIPTION. — Head depressed and broad, body moderately compressed. Dorsal profile rising
evenly but not steeply from tip of snout to origin of dorsal fin, then sloping gently ventrally from there
to end of caudal peduncle. Ventral profile horizontal to origin of anal fin, then sloping dorsally to end
of caudal peduncle. Adipose fin with long base, spanning most of postdorsal distance. In %SL: head
length 26.8, head width 18.2, head depth 13.6, predorsal distance 38.3, preanal length 69.6, prepelvic
length 28.2, prepectoral length 23.8, body depth at anus 15.3, length of caudal peduncle 20.2, depth of
caudal peduncle 9.1, pectoral-spine length 11.0, pectoral-fin length 16.5, dorsal-spine length 15.9,
length of dorsal fin 24.3, length of dorsal-fin base 15.8, pelvic-fin length 7.9, length of anal-fin base
10.7, caudal-fin length 21.4, length of adipose-fin base 30.8, adipose-fin maximum height 4.9,
post-adipose distance 13.8, dorsal to adipose distance 5.0; in %HL: snout length 40.6, interorbital dis-
tance 28.6, eye diameter 23.1, nasal barbel length 36.3, maxillary barbel length 243.i, inner mandibu-
lar barbel length 49.8, outer mandibular barbel length 78.2. Branchiostegal rays 11 (1). Gill rakers
5 + 16 =21 (1). Vertebrae 26 + 24 = 50 (1).

Fin ray counts: dorsal II,7 (1); pectoral I,10 (1); pelvic 1,5 (1); anal iv,7 (1); caudal 1,7,8,1 (1). Dor-
sal-fin origin nearer to tip of snout than to caudal flexure. Dorsal spine stout, without serrations on
posterior edge. Pectoral spine stout, with 1 1 large serrations on posterior edge. Caudal fin forked; dis-
tal margins of upper and lower lobes rounded.

COLOR. — Preserved specimen has the dorsal surfaces of the head and body brown with irregular
dark brown markings forming a variegated pattern; this color fades to a dirty white on the ventral sur-

140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

FIGURE 12. Hemibagrus variegatus, holotype, BMNH 1992.11.16.11; Myanmar: Tenasserim River.

faces. All fins brown, with melanophores on fin rays and interradial membranes; melanophores are
more concentrated in the interradial membranes of the dorsal, pectoral, pelvic and anal fins.

ETYMOLOGY. — From the Latin variegatus, meaning of different colors. In reference to the ir-
regular dark brown markings on the sides of the body.

DISTRIBUTION. — Known only from the Tenasserim River drainage in southern Myanmar
(Fig. 2).

REMARKS. — As mentioned above, H. variegatus is one of the only two known species of south-
ern Asian Hemibagrus (the other being H. imbrifer) which has a long-based adipose fin spanning
nearly all of the postdorsal distance. When compared with other species of Hemibagrus with long adi-
pose-fin bases, H. variegatus has fewer vertebrae (50 vs. 52-60) than H. olyroides and East Asian spe-
cies allied with H. guttatus, and more vertebrae (50 vs. 44-47) than H. baramensis and H. sabanus.

ACKNOWLEDGMENTS

The authors thank the following for permission to examine specimens under their care: William
Saul (ANSP), Darrell Siebert (BMNH), William Eschmeyer (CAS), Sven Kullander (NRM), Douglas
Nelson (UMMZ), Lynne Parenti (USNM) and A. K. Karmakar (ZSI). Maurice Kottelat called our at-
tention to the two specimens that constitute the type series of Hemibagrus imbrifer. Illustrations of
Hemibagrus punctatus and H. variegatus were provided by Kelvin Lim. Funding for this project was
provided by a short-term visitor grant from the Smithsonian Institution to the first author and research
grant RP 960314 from the National University of Singapore to Peter K. L. Ng.

LITERATURE CITED

ANDERSON, J. 1879. Anatomical and zoological researches; comprising an account of the zoological results of the
two expeditions to western Yunnan in 1868 and 1875. Bernard Quaritch, London.

ATAUR RAHMAN, A. K. 1974. Aid to the identification of the mystid catfishes of Bangladesh. Bangladesh Journal
of Zoology 2:1—112.

. 1989. Freshwater fishes of Bangladesh. Zoological Society of Bangladesh, Dhaka. xvii + 364 pp.

BABU RAO, M. AND S. K. CHATTOPADHYAY. 1969. Systematic studies on Mystus spp. (Pisces: Bagridae) of west
Bengal. Journal of the Bengal Natural History Society 35:86—104.

BARMAN, R. P. 1993. Pisces: Freshwater Fishes. Pp. 89-334 in State Fauna Series 5: Fauna of Andhra Pradesh.
Part 1. A. K. Ghosh, ed. Zoological Survey of India, Calcutta.

NG AND FERRARIS: HEMIBAGRUS SPECIES 141

BAILEY, R. M. 1951. The authorship of names proposed in Cuvier and Valenciennes’ “Histoire Naturelle des
Poissons.” Copeia 1951(3):249-251.

BLEEKER, P. 1853. Nalezingen op de ichthyologische fauna van Bengalen en Hindostan. Verhandelingen van het
Bataviaasch Genootschap van Kunsten en Wetenschappen 25: 1—164.

. 1862. Atlas ichthyologique des Indes Orientales Néérlandaises, publié sous les auspices du
Gouvernement colonial néérlandais. Tome II. Siluroides, Chacoides et Héterobranchoides. Lange and Co.,
Amsterdam, 1-112, pls. 49-101.

BLYTH, E. 1860. Report on some fishes received chiefly from the Sitang River and its tributary streams. Journal of
the Asiatic Society (Calcutta) 29:138—174.

BOULENGER, G. A. 1894. Description of a new siluroid fish from Burma. Annals and Magazine of Natural His-
tory, Series 6 14:196.

CUVIER, G. AND A. VALENCIENNES. 1840. Histoire naturelle des poissons. Tome 14. Pitois-Levrault, Paris.
xxiv + 464 pp. [Date of publication corrected from 1839, based on Bailey (1951).]

DAY, F. 1867. On the fishes of the Neilgherry Hills and rivers around their bases. Proceedings of the Zoological
Society of London 1867:281—302.

. 1869. On the fishes of Orissa — Part I. Proceedings of the Zoological Society of London 1869:296—3 10.

. 1871a. On Hamilton Buchanan’s original drawings of fish in the library of the Asiatic Society of Bengal.

Proceedings of the Asiatic Society of Bengal 1871:195—209.

. 1871b. On the freshwater siluroids of India and Burmah. Proceedings of the Zoological Society of Lon-

don 1871:703—721.

. 1877. The fishes of India, being a natural history of the fishes known to inhabit the seas and fresh waters

of India, Burma, and Ceylon. Bernard Quaritch, London. Part 3:369—552, pls. 79-138.

. 1889. The fauna of British India, including Ceylon and Burma. Fishes. Vol. 2. Taylor and Francis, Lon-
don. xiv + 509 pp.

DuTTA, A. K., D. K. KUNDU, AND A. K. KARMAKAR. 1993. Freshwater Fishes. Pp. 1-37 in State Fauna Series |:
Fauna of Orissa. Part 4. A. K. Ghosh, ed. Zoological Survey of India, Calcutta.

ESCHMEYER, W. E., ed. 1998. Catalog of Fishes. California Academy of Sciences, San Francisco. 2905 pp.

GOVIND, B. V. AND K. V. RAJAGOPAL. 1975. Occurrence of a giant Mystus species in Krishna river system.
Matsya 1:79-80.

GRANT, S. A. 1999. A replacement name (nomen nudum) and neotype designation for Hara malabarica Day,
1865, with notes on related species (Siluriformes). Aqua 3:169-174.

GUNTHER, A. 1864. Catalogue of the fishes in the British Museum, volume 5. Catalogue of the Physostomi, con-
taining the families Siluridae, Characinidae, Haplochitonidae, Sternoptychidae, Scopelidae, Stomiatidae in
the collection of the British Museum. British Museum (Natural History), London. xxii + 455 pp.

HAMILTON, F. 1822. An account of the fishes found in the river Ganges and its branches. Privately published, Ed-
inburgh and London. vii + 405, atlas with 39 pls.

Hora, S. L. 1929. An aid to the study of Hamilton Buchanan’s “Gangetic Fishes.’”” Memoirs of the Indian Mu-
seum 9:169—192.

JAYARAM, K. C. 1954. Siluroid fishes of India, Burma and Ceylon XIV. Fishes of the genus Mystus Scopoli. Re-
cords of the Indian Museum 51:527—558.

. 1977. Aid to the identification of the siluroid fishes of India, Burma, Sri Lanka, Pakistan and Bangladesh

1. Bagridae. Records of the Zoological Survey of India, Miscellaneous Publication, Occasional Paper No. 8.

41 pp.

. 1981. The freshwater fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka—a handbook. Zoo-

logical Survey of India, Calcutta. viii + 475 pp.

. 1995. The Krishna River System Bioresources Study. Records of the Zoological Survey of India, Occa-
sional Paper No. 160. 167 pp.

JAYARAM, K. C. AND K. P. SINGH. 1977. On a collection of fish from North Bengal. Records of the Zoological
Survey of India 72:243-275.

JERDON, T. C. 1849. On the fresh-water fishes of southern India. (Continued from p. 149.). Madras Journal of Lit-
erature and Science 15:302—346.

KYAW WIN. 1971. A taxonomy of fishes of Taung-tha-man ‘inn’, upper Burma. Union of Burma Journal of Life
Sciences 4:39-63.

142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 11

MENON, A. G. K. 1977. A check-list of fishes of the Himalayan and Indo-Gangetic plains. Inland Fisheries Soci-
ety of India, Special Publication No. 1. vii + 136 pp.

Mo, T. 1991. Anatomy and systematics of Bagridae (Teleostei) and siluroid phylogeny. Koeltz Scientific Books,
Koenigstein, Germany. vii + 216 pp, 63 pages of figures.

MotTWaANI, M. P., K. C. JAYARAM, AND K. L. SEGHAL. 1962. Fish and fisheries of Brahmaputra river system,
Assam. 1. Fish fauna with observations on their zoogeographical significance. Tropical Ecology 3:17-43.

Noa, P. K. L. AND H. H. NG. 1995. Hemibagrus gracilis, a new species of large riverine catfish (Teleostei:
Bagridae) from Peninsular Malaysia. Raffles Bulletin of Zoology 43:133—142.

Na, H. H. AND W. J. RAINBOTH. 1999. The bagrid catfish genus Hemibagrus (Teleostei: Siluriformes) in central
Indochina with a new species from the Mekong River. Raffles Bulletin of Zoology 47(2):555-576.

QuRESHI, M. R. 1965. Common freshwater fishes of Pakistan. Government Press, Karachi. viii + 61 pp.

RAMAKRISHNIAH, M. 1988. A new bagrid fish of the genus Mystus (Scopoli) from Krishna river system. Matsya
12—13:139-143.

RoBERTS, T. R. 1993. Artisanal fisheries and fish ecology below the great waterfalls of the Mekong River in
southern Laos. Natural History Bulletin of the Siam Society 41:3 1-62.

. 1994. Systematic revision of Asian bagrid catfishes of the genus Mystus sensu stricto, with a new species
from Thailand and Cambodia. Ichthyological Explorations of Freshwaters 5(3):241—256.

ROBERTS, T. R. AND T. J. WARREN. 1994. Observations on fishes and fisheries in southern Laos and northeastern
Cambodia, October 1993—February 1994. Natural History Bulletin of the Siam Society 42:87-115.

ROSSEL, F. 1964. Welse (Siluroidea), gesammelt von de deutschen Indien-Expedition 1955/58. Mitteilungen aus
dem Hamburgischen Zoologischen Museum und Institut 61:145—158.

SEN, T. K. 1985. The Fish Fauna of Assam and the Neighbouring North-eastern States of India. Occasional Paper
of the Zoological Survey of India No. 64. 216 pp.

. 1992. Freshwater fish. Pp. 101—242 in State Fauna Series 3: Fauna of West Bengal. Part 2. Reptilia,
Amphibia, Fishes, Hemichordata and Archaeozoology. A. K. Ghosh, ed. Zoological Survey of India, Cal-
cutta.

SHAW, G. E. AND E. O. SHEBBEARE. 1937. The fishes of northern Bengal. Journal of the Royal Asiatic Society,
Science 3:1—137, 6 pls.

SHRESTHA, J. 1994. Fishes, fishing implements and methods of Nepal. Smt. M. D. Gupta, Lashkar. Iv + 150 pp.

SINGH, D. F. AND G. M. YAZDANI. 1993. Ichthyofauna of Konkan region of Maharashtra (India). Records of the
Zoological Survey of India, Occasional Paper No. 145. 46 pp.

SRIVASTAVA, G. J. 1968. Fishes of eastern Uttar Pradesh. Vishwavidyalaya Prakashan, Varanasi. xxii + 163 pp.

TALWAR, P. K. AND A. G. JHINGRAN. 1991. Inland fishes of India and adjacent countries. Oxford and IBH Pub-
lishing Company, New Delhi, 2 vols., 1158 pp.

TINT HLIANG. 1971. A classified list of fishes of Burma. Union of Burma Journal of Life Sciences 4:507—526.

UKKATAWEWAT, S. AND C. VIDTHAYANON. 1998. Fishes of the Salween River basin, Thailand. Aquatic Natural
Resources Museum Technical Paper 2:1—89.

VINCIGUERRA, D. 1890. Viaggo di Leonardo Fea in Birmania e regioni vicini. XXIV. Pesci. Annali del Museo
Civico de Storia Naturale de Genova (serie 2a) 9:129—362, pls. 7-11.

VISHWANATH, W. AND H. T. SINGH. 1986. First record of the bagrid catfish Mystus microphthalmus from India.
Japanese Journal of Ichthyology 33:197—199.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 12, pp. 143-158, figs. 1-3, 1 table. December 7, 2000

Chromosome Numbers of South African Acanthaceae

by
Thomas F. Daniel

Department of Botany, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and
Kevin Balkwill and Mandy-Jane Balkwill

C. E. Moss Herbarium
Department of Animal, Plant, and Environmental Sciences
University of the Witwatersrand
Private Bag X3, WITS 2050, South Africa

Original meiotic chromosome counts are presented for 19 species in 14 genera of
Acanthaceae from South Africa. These counts represent the initial reports of chromosome
numbers from southern African Acanthaceae. Chromosome numbers of 13 species repre-
senting 11 genera are reported for the first time. The counts in Aulojusticia, Duvernoia, and
Metarungia are the first for these genera. Counts for five species confirm numbers previ-
ously reported for them based on plants from other regions. A new chromosome number is
reported in Justicia (n = 26 in J. petiolaris). Systematic implications of these chromosome
counts are addressed.

The Acanthaceae are a large (ca. 4000 species in some 230 genera) pantropical family with major
concentrations of species in southeastern mainland Asia, insular Malesia, the Indian subcontinent,
Madagascar, tropical Africa, Brazil, Andean South America, and Mexico-Central America. Knowl-
edge of chromosome numbers among Acanthaceae has proven useful in resolving generic positions of
problematic species, reassessing phylogenetic relationships among subfamilial taxa, and understand-
ing morphological variation (e.g., Ensermu Kelbessa 1990; Daniel and Chuang 1993; McDade et al.
in press). A major problem with using chromosome number data in the study of systematic and evolu-
tionary relationships among Acanthaceae is that numbers remain unknown for the vast majority of
both genera and species. Daniel (2000) noted that chromosome numbers had been reported for only
29% of the genera and less than 12% of the species of Acanthaceae. Similarly, chromosome numbers
remain undetermined for five of the 12 tribes recognized by Bremekamp (1965) in the family:
Haselhoffieae, Louteridieae, Rhombochlamydeae, Stenandriopsideae, and Whitfieldieae. Another
problem is the lack of, or poor sampling of, species from regions rich in Acanthaceae but
underrepresented by chromosome counts of them. One such region, as identified by Daniel (2000), is
southern A frica. This region, comprising Namibia, Botswana, South Africa, Lesotho, and Swaziland,
has an acanthaceous flora of some 341 species (Welman 1993). Chromosome numbers have been re-
ported previously for 28 of these (see Appendix), but none of these reports was based on collections
from southern A frica. This study of chromosome numbers in selected Acanthaceae from South Africa
is our first attempt to provide this systematically useful information for taxa from this region.

i - iia

143 r nied

FEB 95 200]

144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

METHODS

In February and March of 1999, floral buds and herbarium vouchers of South African
Acanthaceae were collected from their native habitats and from plants cultivated in gardens. Buds
were fixed in absolute ethanol:glacial acetic acid (3:1) for 24 hours and subsequently washed and
stored in ethanol (70%) until analyzed. Anthers were macerated in ferric acetocarmine (1%) on a mi-
croscope slide, squashed under a coverslip by hand, and studied under oil immersion on a phase con-
trast microscope at a magnification of 1000x. Counts were made from microsporocytes in various
stages of meiosis. Preparations from which counts were obtained were recorded with camera lucida
drawings. Voucher specimens are deposited at CAS and J. The camera lucida drawings are attached to
the vouchers at CAS.

In the following discussion, all previously published chromosome counts are listed as n (gametic,
haploid, meiotic) numbers irrespective of whether they were originally reported as gametophytic or
sporophytic numbers. Voucher specimens, if they exist, documenting previous counts by other work-
ers have not been examined.

RESULTS

Chromosome numbers determined for 19 species representing 14 genera of Acanthaceae from
South Africa are summarized in Table | and illustrated in Figures 1-3. Chromosome numbers of 13 of
these taxa are reported for the first time. The counts for Aulojusticia, Duvernoia, and Metarungia rep-
resent the first reports for these genera. Counts for five species (Asystasia gangetica, Hypoestes
aristata, H. forskaolii, Justicia betonica, and Rhinacanthus gracilis) agree with some or all numbers
previously reported for them from different sources. The chromosome number reported here for
Justicia odora is the first non-approximate number reported for the species. The count of n = 26 for J.
petiolaris is the first report of this number in the genus.

DISCUSSION

Asystasia Blume. Fifty or more species are recognized in this genus, which is native to the Old
World. Eight species are known from southern Africa (Welman 1993). Daniel (2000) summarized
previous reports of chromosome numbers in both A. gangetica and in the genus. Although a diversity
of meiotic numbers, including 14, 22, 24 and 25, has been reported for A. gangetica, counts of n = 13
and n = 26 are much more common. A meiotic number of 13 is also the most frequently reported chro-
mosome number among other species in the genus.

Asystasia gangetica has a broad distribution, occurring indigenously in southern and tropical Af-
rica and in the Indian subcontinent. It has become naturalized in several other tropical regions (e.g.,
Hawaii, Java). Previous counts (including those reported for A. coromandeliana Nees; see summary
of counts in Daniel 2000) have come from plants in cultivation (without provenance data; e.g., Grant
1955), from the Indian subcontinent (e.g., Valsala Devi and Mathew 1982), from tropical west Africa
(e.g., Gadella 1977), and from plants naturalized outside of the native range of the species (e.g., Daniel
2000). Our count of n = 13 for a southern African representative of the species reveals a continuity of
chromosome number with plants from other portions of the species’ range. Plants from which our
count was determined would appear to be diploid within the species. As indicated by Daniel (2000), a
basic number of x = 13 appears likely for the genus.

Aulojusticia Lindau. Our count of n = 40 for A. linifolia is the first chromosome count for this ge-
nus. Aulojusticia has been variably treated by students of Acanthaceae: Dyer (1975) recognized it asa
unispecific genus endemic to northeastern South Africa; Graham (1988) included it within her broad

DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 145

TABLE |. Meiotic chromosome counts of South African Acanthaceae. Note: Counts for Duvernoia
aconitiflora, Justicia petiolaris, Metarungia longistrobus, Pseuderanthemum hildebrandtii, and Ruspolia
hypocrateriformis were obtained from plants in cultivation; * indicates first counts for taxa.

Chromosome Locality
number (7)

Collection
(province) number

Taxon

(Daniel et al.)

Asystasia gangetica (L.) T. Anderson 13 KwaZulu-Natal 9348
Aulojusticia linifolia Lindau 40* Mpumalanga 9388
Barleria senensis Klotzsch 16* Mpumalanga 9375
Crabbea angustifolia Nees 2h Mpumalanga 9370
Dicliptera heterostegia Presl. ex Nees ES* KwaZulu-Natal 9328
Dicliptera magaliesbergensis K. Balkwill 13* Gauteng 9350
Duvernoia aconitiflora A. Meeuse Wh Gauteng 9361
Hypoestes aristata (Vahl) Sol. ex R. & S. 30 KwaZulu-Natal 9351
Hypoestes forskaolii (Vahl) R.Br. 15 Gauteng 9358
Isoglossa hypoestiflora Lindau Ihe KwaZulu-Natal 9341
Isoglossa ovata (Nees) Lindau i7* KwaZulu-Natal 9336
Justicia betonica L. 17 KwaZulu-Natal 9330
Justicia betonica L. i, KwaZulu-Natal 9334
Justicia odora (Forssk.) Lam. 14 North-West 9364
Justicia petiolaris (Nees) T. Anderson 26* Mpumalanga 9387
Metarungia longistrobus (C. B. Cl.) Baden 14* Gauteng 9355
Pseuderanthemum hildebrandtii Lindau Zs Mpumalanga 9394
Rhinacanthus gracilis Klotzsch 15 KwaZulu-Natal 9340
Rhinacanthus gracilis Klotzsch 15 KwaZulu-Natal 9328
Ruspolia hypocrateriformis (Vahl) a Gauteng 9559
Milne-Redh.
Ruttya ovata Harv. 21* KwaZulu-Natal 9338

circumscription of Justicia; and Immelman (1995a) treated it as Siphonoglossa linifolia (Lindau)
C. B. Clarke. Siphonoglossa, which is based on an American type, is conspecific with Justicia (Gra-
ham 1988, Daniel 1995). Molecular studies that seek to improve our understanding of Justicia and al-
lied genera in Africa are currently underway. Aulojusticia undoubtedly falls within the morphological
circumscription of Justicia as delimited by Graham. If treated in that genus, the chromosome number
here reported for A. linifolia would be the highest number so far known in Justicia; the highest number
previously reported is n = 34 (Daniel 2000). Also, if treated in Justicia, which appears to have a basic
number of x = 7 (see below), then this species likely would have been derived from a hexaploid ances-
tor.

Barleria L. Barleria is a pantropical genus of perennial herbs and shrubs comprising some 300
species. The majority of species are African and 69 occur in southern Africa (Balkwill and Balkwill
1997). Daniel and Chuang (1989, 1998) summarized previously reported chromosome numbers in
Barleria. They noted the prevalence of n = 20 among species of the genus and the likelihood of x = 20
as a basic number in the genus.

Our count of n = 16 for Barleria senensis is the first count for this species, which is native to
southern and tropical Africa. Previously, this number has been reported in the genus once for B.
cristata L. (Datta and Maiti 1970, as “B. cristata var. dichotoma’’). There are many counts for this spe-
cies and most of them are n = 20 (Daniel and Chuang 1989). Because these two species are treated in

146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

different subgenera of Barleria (Balkwill and Balkwill 1997), a common chromosome number be-
tween them is doubtfully due to common ancestry.

Barleria was treated in tribe Barlerieae by Lindau (1895) and in tribe Ruellieae, subtribe
Barleriinae by Bremekamp (1965) and Balkwill and Getliffe Norris (1988). Based on DNA sequence
data (Hedrén et al. 1995; Scotland et al. 1995; McDade and Moody 1999), Crabbea and Lepidagathis
would appear to be closely related to Barleria. A chromosome number of n = 21 has been reported in
species from each of these three genera (Chuang etal. 1963; Grant 1964; De 1966; Daniel in press, see
below) suggesting a possible symplesiomorphic number among them. Additional counts of species in
Barleria are needed in order to ascertain whether knowledge of chromosome numbers will help to re-
solve infrageneric relationships.

Crabbea Harv. Two previous chromosome counts have been reported for this African genus of
about 12 species, seven of which occur in southern Africa (Welman 1993). Our count of n = 21 for the
South African endemic, C. angustifolia, is the first report of a chromosome number in the species. It
agrees with a previous count for C. velutina S. Moore (Grant 1964, as C. reticulata C. B. Clarke).
Renard et al. (1983) reported n = ca. 14 for C. velutina.

Based on the few counts for species of Crabbea, a basic number of x = 21 is tentatively proposed
for the genus. Lindau (1895) included Crabbea in his tribe Barlerieae and the genus would be included
in Bremekamp’s (1965) tribe Ruellieae, subtribe Barleriinae. As noted above under Barleria, this
number is known in the related genera Barleria and Lepidagathis. Balkwill and Getliffe Norris (1988)
advocated the placement of Crabbea in the Ruellieae, Ruelliinae. Among genera of Ruelliinae for
which chromosome numbers are known, n = 21 has been reported in several species of Eranthemum
L. (e.g., Kaur 1970; Govindarajan and Subramanian 1985; Mangenot and Mangenot 1958, 1962).

Dicliptera Juss. Chromosome numbers have been reported for 22 of the more than 100 species of
perennial herbs and shrubs in this pantropical genus. Fifteen of these species are from the New World
and seven are native to the Old World. Counts for all New World species are n = 40 (or n = ca. 40;
Daniel 2000, Daniel and Chuang 1993, Piovano and Bernardello 1991) whereas previous counts of
species from the Old World comprise n = 10, 13, 15, 24, and 26 (with n = 13 being most frequently re-
ported; Daniel and Chuang 1993). The species of Dicliptera from the Old World that were studied are
all indigenous to Asia and comprise: D. bupleuroides Nees, D. cuneata Nees, D. elegans W. W. Smith
[Kaur (1970) identified this taxon as “Dicliptera elegans Dalz.,” and neither the source of the plant
nor a voucher was cited; Dalziel did not publish a species with this name and whether the plant studied
by Kaur is actually the Chinese species described by Smith or an Indian taxon is not known], D.
leonotis Dalziel ex C. B. Clarke, D. parvibracteata Nees, D. roxburghiana Nees, and D. verticillata
(Forsk.) C. Chr. Although two of these species (1.e., D. leonotis and D. verticillata) also occur in tropi-
cal Africa, chromosome numbers for all of the Old World species of Dicliptera were apparently deter-
mined from Asian populations. Our counts would therefore appear to be the first from African plants,
and the first for species of Dicliptera native to southern Africa. Twelve species of Dicliptera are
known to occur in southern Africa (Balkwill et al. 1996). The count of n=13 for D.
magaliesbergensis agrees with the majority of previous counts for Old World species. The count of
n= 15 for D. heterostegia agrees with a count by Kaur (1970) for the widespread Paleotropical spe-
cies D. verticillata.

Daniel et al. (1990) noted that the difference in ploidal level between species of Dicliptera in the
Old and New Worlds suggests a major geographical division in the genus. Recent phylogenetic stud-
ies of the Justicieae that included Paleotropical and Neotropical species of Dicliptera indicate that the
genus is monophyletic and is related to Hypoestes and Peristrophe in subtribe Diclipterinae (McDade
et al. in press).

DANIEL, BALKWILL, AnD BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 147

i) oe =
& S
é o “FR “e
* o* o-
w 0 ws)
O4,%% 0 ° ® 6? e
a* %
ae Bae 38
é “Me re sal
| oe @
a ea
a 2,6 b
10 um

@ e a @
“ae “ ® fet ts
r) | CP w™ e
9? ms ® te \ f ed
eS Oe on
eo re

FIGURE 1. Camera-lucida drawings of meiotic chromosome preparations. a. Justicia petiolaris (Daniel et al. 9387), n = 26
(telophase I). b. Ruttya ovata (Daniel et al. 9338), n =21 (metaphase I). c. Dicliptera heterostegia (Daniel et al. 9329),n=15
(diakinesis I). d. Duvernoia aconitiflora (Daniel et al. 9361 ), n= 17 (metaphase I). e. Jsoglossa ovata (Daniel et al. 9336),
n= 17 (telophase I). f. Pseuderanthemum hildebrandtii (Daniel et al. 9394), n = 21 (diakenesis 1).

Piovano and Bernardello (1991) noted that, based on chromosome numbers so far reported in
Dicliptera, x = 10 is likely the basic number of the genus. The occurrence of n = 15 in species of all
genera of Diclipterinae, however, suggests that x = 15 is symplesiomorphic for both the subtribe and
Dicliptera (Daniel and Chuang 1993, McDade et al. in press).

148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

Duvernoia E. Mey. ex Nees. This genus of two species that are endemic to eastern South Africa
and Mozambique is sometimes treated within Justicia (Graham 1988; Brummitt 1992). Bremekamp
(1939) argued for the recognition of Duvernoia, primarily on the basis of its pollen, and treated at least
one species from the New World in it. Our count of n = 17 for D. aconitiflora is the first report of a
chromosome number for both the genus and the species. Manning and Getliffe Norris (1995) treated
the genus as distinct from others in the tribe Justicieae, subtribe Justiciinae on the basis of its
pedunculate inflorescences with small bracts and campanulate calyces. It remains to be determined
whether these characteristics are sufficient to delimit the two species sometimes treated in Duvernoia
from other Justiciinae on a global basis. A meiotic complement of 17 is known, but infrequently re-
ported, in Justicia; it has been noted in four species, J. adhatoda L. (e.g., Daniel and Chuang 1998), J.
betonica L. (e.g., Daniel and Chuang 1998 and see below), J. carnea Lindl. (Daniel and Chuang
1989), and J. trinervia Vahl (e.g., Krishnappa and Ranganath 1982), representing three sections of the
genus (Graham 1988). It is perhaps revealing that J. adhatoda is the type of one of these, section
Vasiaca Lindau, in which Graham (1988) placed the other species of Duvernoia, D. adhatodoides E.
Mey ex Nees.

Hypoestes Sol. ex R.Br. Hypoestes consists of about 70 species of herbs and shrubs from Africa,
eastern Asia, Malesia, and Australia. Three species occur in southern Africa (Balkwill and Getliffe
Norris 1985). Our count of n = 15 for H. forskaolii, a species occurring in Africa and the Arabian pen-
insula, agrees with a previously reported chromosome number for this species (Podlech 1986) and
with most numbers reported for other species in the genus (Daniel and Chuang 1998). Our count of
n = 30 for H. aristata, a species native to tropical and southern A frica, agrees with a previous count for
this species based on cultivated plants (Daniel and Chuang 1998). This is the only known species of
Hypoestes with a chromosome number other than n = 15. If a basic number of x = 15 is accepted for
Hypoestes then H. aristata would represent a tetraploid species in the genus. A haploid chromosome
number of 15 is also known in Dicliptera, Peristrophe, and Rhinacanthus (see below), all southern
African relatives of Hypoestes in tribe Justicieae, subtribe Diclipterinae. Our collection of H. aristata
is referable to var. alba K. Balkwill.

Isoglossa Oerst. Isoglossa consists of about 60 species native to the Old Worid. Welman (1993)
listed 15 species as occurring in southern Africa. Our counts of n = 17 for two of these, /. hypoestiflora
(southern and tropical Africa) and /. ovata (southern Africa), agree with the sole previous report of a
chromosome number in the genus. Daniel and Chuang (1998) reported n = 17 for the tropical eastern
African species /. grandiflora C. B. Clarke.

Tsoglossa was treated by Bremekamp in tribe Justicieae, subtribe Rhytiglossinae
(= Isoglossinae). McDade et al. (in press) demonstrated that /sog/ossa is part of a “core” Isoglossinae
that is strongly supported as monophyletic. Chromosome numbers have been determined for only two
other genera in this subtribe; Daniel (1999) reported n = 18 for both Stenostephanus Nees (including
Habracanthus Nees and Hansteinia Oerst.) and Razisea Oerst. Although a basic number of x = 17 ap-
pears likely for /soglossa, and may represent dysploid evolution from an ancestor with x = 18, a
well-substantiated basic number for the subtribe remains to be determined.

Justicia L. Justicia is the largest genus of Acanthaceae with estimates of about 700 species occur-
ring worldwide (McDade et al. in press). Twenty-two species were recognized by Immelman (1995b)
in the treatment of Acanthaceae: Justicieae for the Flora of Southern Africa. Several other genera
from the region that are recognized in the Flora of Southern Africa (1.e., Adhatoda Mill., Aulojusticia,
Duvernoia, Siphonoglossa Oerst.) are sometimes included in Justicia (Graham 1988, Brummitt
1992) as well. Daniel (2000) noted the presence, frequency, and distribution of 21 chromosome num-
bers reported for 93 species in the genus. He noted the prevalence of n = 14 throughout Justicia and a

DANIEL, BALKWILL, AND BALK WILL: ACANTHACEOUS CHROMOSOME COUNTS 149

me a o®
eo, *
%& V a ct
4 — °
oe &
ps “te L
a
ee
& ot Pg
)
*
f * ge
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4
oN is o << en
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10 um at. 4 $
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FIGURE 2. Camera-lucida drawings of meiotic chromosome preparations. a. Justicia betonica (Daniel et al. 9334),n=17
(metaphase I). b. Jsoglossa hypoestiflora (Daniel et al. 9341), n = 17 (diakenesis I). c. Barleria senensis (Daniel et al. 93 75),
n= 16 (telophase 1). d. Asystasia gangetica (Daniel et al. 9348), n = 13 (metaphase 1). e. A ulojusticia linifolia (Daniel et al.
9388),n = 40 (metaphase I). f. Justicia odora (Daniel et al. 9364), n = 14 (diakenesis I). g. Crabbea angustifolia (Daniel et al.
9370), n = 21 (early metaphase 1).

150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

probable basic number of x = 7 for the genus. Within Justicia several sections are characterized by a
diversity of chromosome numbers whereas others are homogeneous in the counts reported (Daniel
2000).

Our counts for three species of southern African Justicia illustrate the diversity of numbers re-
ported in the genus. The counts of m = 17 for two collections of J. betonica agree with the majority of
previous counts for this native of Africa and the Indian subcontinent (Daniel 2000, Daniel and Chuang
1998). Hedrén (1989) reported n = ca. 13 from a Tanzanian collection of the African and Arabian spe-
cies, J. odora. Our count for this species, based ona collection from South Africa, ism = 14. Thisnum-
ber has been reported previously in 10 of the 13 sections of Justicia recognized by Graham (1988) in
which at least one chromosome number has been reported. This is the first report of n = 14 in section
Harnieria (Solms) Benth., however. Our count of n = 26 inJ. petiolaris, a species occurring in eastern
South Africa and Swaziland, represents the first report of a chromosome number for the species. It is
also the first report of this chromosome number in the genus. Graham (1988) treated J. petiolaris in
sect. Tyloglossa (Hochst.) Lindau. The only other chromosome number known among the species in-
cluded in this section by Graham is n = 13, which has been reported several times (e.g., Mangenot and
Mangenot 1962, Podlech 1986) in J. flava (Vahl) Vahl. Our collection of /. petiolaris pertains to sub-
species petiolaris. Chromosome numbers reported here for South African species of Justicia reflect
the putative polyploid and dysploid evolution seen for the genus in other regions (Daniel 2000). Given
the large number of species of Justicia, its worldwide distribution, and the diversity of chromosome
numbers already reported within it, additional cytological studies of the genus are highly desirable.

Metarungia Baden. This genus comprises three species occurring in eastern and southern Africa
(Baden 1981, 1984). Our count of n = 14 for M. longistrobus, a species native to South Africa, Swazi-
land, and Mozambique, is the first report of a chromosome number in the genus. Metarungia is closely
related to Rungia Nees in the tribe Justicieae and these genera are sister to a clade that includes Justicia
(McDade et al. in press). Numerous counts have been reported for each of four species of Rungia.
Counts for three of these (i.e., R. Jaeta C. B. Clarke, R. parviflora Nees, and R. pectinata (L.) Nees) are
mostly n = 13 or n= 26. Most counts for Rungia repens Nees are n= 10. Although n = 14 is not
known in Rungia, this is the most commonly reported number in Justicia (Daniel 2000).

Pseuderanthemum Radlk. Pseuderanthemum consists of about 60 species of perennial herbs and
shrubs occurring in both the Old and New Worlds. Two species are known from southern Africa, P.
subviscosum (C. B. Clarke) Stapf and P. hildebrandtii (Welman 1993; Edwards and Harrison 1998).
Our count of = 21 for P. hildebrandtii, which is native to southern Africa and tropical east Africa, is
the first report of a chromosome number for this species and for an African species of the genus. Chro-
mosome numbers have been reported for seven other species of Pseuderanthemum native to America,
southern Asia, and the Pacific Islands (Daniel and Chuang 1989, 1998; Daniel 2000). Six of these
have meiotic complements of 21. Kaur (1969) reported n = 30 for the Fijian species, P. laxiflorum (A.
Gray) F. T. Hubb. As noted by Daniel and Chuang (1998), a basic number of x = 21 appears likely for
this genus as well as for several of its relatives in the Justicieae. McDade et al. (in press) identified a
lineage of Justicieae, including Pseuderanthemum, characterized by an androecium of two stamens
and two staminodes and by x=21. New World relatives include Chileranthemum Oertst.,
Odontonema Nees, and Oplonia Raf. Old World relatives include Ruspolia and Ruttya.

Rhinacanthus Nees. Rhinacanthus comprises about 20 species occurring in Africa, Madagascar,
and Asia. Three species are known in southern Africa (Balkwill 1995). One of these, R. gracilis, is
sometimes cultivated. Daniel and Chuang (1998) reported a count of n = 15 from cultivated materials
of this species. Our count of n = 15 from two wild-collected plants of the same species confirms the
previous count. All other counts reported for species of Rhinacanthus are likewise n = 15 (Daniel and

DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS ite

r Pans °

d a % p e &

ad me a
“pe te
& SG ES o

® 4

s <a

f e

FIGuRE 3. Camera-lucida drawings of meiotic chromosome preparations. a. Hypoestes aristata (Daniel et al. 9351), n = 30
(telophase II, only half of cell shown). b. Rhinacanthus gracilis (Daniel 9340), n=15 (metaphase I). c. Dicliptera
magaliesbergensis (Daniel et al. 9357), n= 13 (telophase I). d. Ruspolia hypocrateriformis (Daniel et al. 9359), n=21
(diakenesis I). e. Hypoestes forskaolii (Daniel et al. 9358), n = 15 (diakenesis I). f. Metarungia longistrobus (Daniel et al.

9355), n = 14 (telophase I).

152 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

Chuang 1998). Thus, a basic number of x = 15 can be advanced for the genus. Based on morphological
characteristics Rhinacanthus would appear to be best treated in tribe Justicieae, subtribe Justiciinae
(Balkwill and Getliffe Norris 1988), but molecular sequence data (McDade et al. in press) place it in
subtribe Diclipterinae with strong support. Cytological data corroborate this latter placement. Other
Diclipterinae (i.e., Dicliptera, Hypoestes, and Peristrophe) also have probable basic chromosome
numbers of 15 (see above).

Nomenclatural and taxonomic confusion surrounding R. gracilis was discussed by Balkwill
(1995), who recognized two varieties of the species in southern Africa. Plants used for our counts per-
tain to R. gracilis var. latilabris K. Balkwill. The voucher specimen of Daniel and Chuang’s (1998)
count pertains to R. gracilis var. gracilis.

Ruspolia Lindau. Six species of Ruspolia have been described from Africa and Madagascar and
three of them occur in southern Africa (Welman 1993). Our count of n = 21 for R. hypocrateriformis,
a species native to western tropical Africa and South Africa, is the first report of a chromosome num-
ber for the species. It agrees with previous counts of n= 21 for R. seticalyx (C.B. Clarke)
Milne-Redh. (Daniel and Chuang 1998). This same chromosome number has been reported in several
relatives of Ruspolia in the Justicieae (see above under Pseuderanthemum). Plants from South Africa
are commonly treated as R. hypocrateriformis var. australis Milne-Redh. (e.g., Heine 1963).

Ruttya Harv. Ruttya comprises three species native to Africa. Only one of these, R. ovata, occurs
in southern A frica where it is found in South Africa and Swaziland. Our count of = 21 for the species
agrees with the only previous report of a chromosome number in the genus. Daniel and Chuang (1998)
reported this number for R. fruticosa Lindau, a native of eastern tropical Africa, based on cultivated
plants. The affinities of Ruttya are with Ruspolia and other genera of tribe Justicieae with an androe-
cium of two stamens and two staminodes and x = 21 (see above under Pseuderanthemum). Morpho-
logical distinctions among Ruttya, Ruspolia, and Pseuderanthemum were noted by Daniel and
Chuang (1998).

CONCLUSIONS

As shown by McDade et al. (in press), knowledge of chromosome numbers can be useful for sug-
gesting or corroborating phylogenetic relationships among Acanthaceae. Even in this small sampling
of taxa from South Africa, the importance of knowledge of chromosome numbers among
Acanthaceae is evident. A continuity of chromosome numbers between southern African endemics
and congeners from other regions provides additional evidence for several previously proposed
suprageneric relationships based on morphology and molecular markers (e.g.,
Hypoestes-Dicliptera-Peristrophe and Pseuderanthemum-Ruttya-Ruspolia). However, the chromo-
some numbers reported for Rhinacanthus support molecular rather than morphological data in plac-
ing this genus in the Diclipterinae rather than in the Justiciinae. Based on the treatment of Justiciinae
in the Flora of Southern Africa (Baden et al. 1995), Aulojusticia and Duvernoia are recognized here.
Using morphological characteristics, both might be readily accommodated in a broadly interpreted
Justicia. However, their respective chromosome numbers are either unknown or uncommon in that
genus and their systematic affinities based on molecular markers are currently being studied.

It will be useful to obtain additional chromosome numbers among Acanthaceae from southern
Africa by sampling more taxa from South Africa as well as from other countries in the region. For ex-
ample, chromosome numbers remain unknown for the following acanthaceous genera that occur in
southern Africa: Acanthopsis Harv., Anisotes Nees, Chaetacanthus Nees, Chorisochora Vollesen,
Duosperma Dayton, Glossochilus Nees, Megalochlamys Lindau, Neuracanthus Nees, and
Ruelliopsis C. B. Clarke. Also, no chromosome counts have been determined for Acanthaceae from

DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 153

any southern African country except South Africa. An increase in the number of taxa for which chro-
mosome numbers are known would help to identify basic numbers for additional genera and assist ef-
forts to understand phylogenetic relationships within the family.

ACKNOWLEDGMENTS

We are grateful to Fei Mei Chuang for technical assistance with the chromosome counts, R. K.
Brummitt and M. Butterwick for assistance in the field, J. and S. Williamson for logistical assistance
and for allowing us to collect plants in their garden, and the Witwatersrand National Botanical Garden
and the Lowveld National Botanical Garden for permitting us to collect buds and vouchers of
Acanthaceae. Daniel’s field work in South Africa was funded by the In-House Research Fund of the
California Academy of Sciences and the field work of K. and M.-J. Balkwill was funded by the Na-
tional Research Foundation of South Africa and the University of the Witwatersrand’s Herbarium
Programme.

LITERATURE CITED

BADEN, C. 1981. The genus Macrorungia (Acanthaceae), a taxonomic revision. Nordic J. Bot. 1:143-153.

. 1984. Metarungia, a valid name for Macrorungia auctt. (Acanthaceae). Kew Bull. 39:638.

BADEN, C., K. BALKWILL, F. M. GETLIFFE NorRIS, K. L. IMMELMAN, J. C. MANNING, AND J. MUNDAY. 1995.
Acanthaceae, Fascicle 1: Justiciinae. /n Flora of southern Africa, O. A. Leistner, ed. 30(3, 1):1—71.

BALKWILL, K. 1995. 3. Rhinacanthus. In Flora of southern Africa, O. A. Leistner, ed. 30(3, 1):11—14.

BALKWILL, M.-J. AND K. BALKWILL. 1997. Delimitation and infra-generic classification of Barleria
(Acanthaceae). Kew Bull. 52:535—-573.

BALKWILL, K. AND F. GETLIFFE NORRIS. 1985. Taxonomic studies in the Acanthaceae; the genus Hypoestes
Soland. ex R. Br. in southern Africa. S. Afr. J. Bot. 51:133-144.

. 1988. Classification of the Acanthaceae: a southern African perspective. Monogr. Syst. Bot. Missouri
Bot. Gard. 25:503-516.

BALKWILL, K., F. GETLIFFE NORRIS, AND M.-J. BALKWILL. 1996. Systematic studies in the Acanthaceae;
Dicliptera in southern Africa. Kew Bull. 51:1-61.

BREMEKAMP, C. E. B. 1939. Acanthaceae. Jn LIX—Contributions to the Flora of Tropical America: XLIII, N.
Y. Sandwith et al., eds. Kew Bull. 1939:545-563.

. 1965. Delimitation and subdivision of the Acanthaceae. Bull. Bot. Surv. India 7:21—30.

BRUMMITT, R. K. 1992. Vascular plant families and genera. Royal Botanic Gardens, Kew.

CHUANG, T. I., C. Y. CHAO, W. W. L. Hu, AND S. C. KWAN. 1963. Chromosome numbers of the vascular plants
of Taiwan. I. Taiwania 8:51—66.

DANIEL, T. F. 1995. Acanthaceae. /n Flora of Chiapas, Part 4, D. E. Breedlove, ed. California Academy of Sci-
ences, San Francisco. 158 pp.

. 1999. Revision of Stenostephanus (Acanthaceae) in Mexico. Contr. Univ. Michigan Herb. 22:47-93.

. 2000. Additional chromosome numbers of American Acanthaceae. Syst. Bot. 25:15—25.

. In press. Chromosome numbers of some Acanthaceae from Papua New Guinea. Austrobaileya.

DANIEL, T. F. AND T. I. CHUANG. 1989. Chromosome numbers of some cultivated Acanthaceae. Baileya
23:86-93.

. 1993. Chromosome numbers of New World Acanthaceae. Syst. Bot. 18:283-289.

. 1998. Chromosome numbers of cultivated Acanthaceae and systematic implications. Pp. 309-330 in
Diversity and taxonomy of tropical flowering plants, P. Mathew and M. Sivadasan, eds. Mentor Books,
Calicut.

DANIEL, T. F., T. I. CHUANG, AND M. A. BAKER. 1990. Chromosome numbers of American Acanthaceae. Syst.
Bot. 15:13-25.

DaTTA, P. C. AND R. K. MATAI. 1970. Relationships of Justicieae (Acanthaceae) based on cytology. Genetica
41:437-4S0.

154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

De, A. 1966. Cytological, anatomical and palynological studies as an aid in tracing affinity and phylogeny in the
family Acanthaceae. I. Cytological Studies. Trans. Bose Res. Inst. 29:139-175.

Dyer, R. A. 1975. The genera of the southern African flowering plants, vol. 1. Department of Agricultural Tech-
nical Services, Pretoria.

EDWARDS, T. J. AND E. HARRISON. 1998. New records from KwaZulu-Natal, South Africa. Bothalia 28:187—190.

ENSERMU KELBESSA. 1990. Justicia sect. Ansellia (Acanthaceae). Acta Universitatis Upsaliensis Symbolae
Botanicae Upsalienses 29(2):1—96.

GADELLA, T. W. J. 1977. In IOPB chromosome number reports LVI. Taxon 26:259-261.

GOVINDARAJAN, T. AND D. SUBRAMANIAN. 1985. Karyomorphological studies in South Indian Acanthaceae.
Cytologia 50:473-482.

GRAHAM, V. A. W. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bull.
43:551-624.

GRANT, W. F. 1955. A cytogenetic study in the Acanthaceae. Brittonia 8:121—149.

. 1964. Jn IOPB chromosome number reports. I. Taxon 13:108.

HEDREN, M. 1989. Justicia sect. Harnieria (Acanthaceae) in tropical Africa. Acta Universitatis Upsaliensis
Symbolae Botanicae Upsalienses 29(1):1—141.

HEDREN, M., M. W. CHASE, AND R. G. OLMSTEAD. 1995. Relationships in the Acanthaceae and related families
as suggested by cladistic analysis of rbcL nucleotide sequences. Pl. Syst. Evol. 194:93—109.

HEINE, H. 1963. Acanthaceae. Pp. 391-432 in Flora of west tropical Africa 2(ed. 2), F. N. Hepper, ed.

IMMELMAN, K. L. 1995a. 2. Siphonoglossa. In Flora of southern Africa, O. A. Leistner, ed. 30(3, 1):6—10.

. 1995b. 5. Justicia. In Flora of southern Africa, O. A. Leistner, ed. 30(3, 1):18-46.

KAUR, J. 1969. Chromosome numbers in Acanthaceae—IV. Sci. and Cult. 35:61-63.

. 1970. Chromosome numbers in Acanthaceae—V. Sci. and Cult. 36:103—106.

KRISHNAPPA, D. G. AND R. M. RANGANATH. 1982. /n IOPB Chromosome number reports LXXV. Taxon
31:364-36S5.

LINDAU, G. 1895. Acanthaceae. /n Die natiirlichen Pflazenfamilien, A. Engler and K. Prantl, eds. 4(3b):274-354.
MANGENOT, S. AND G. MANGENOT. 1958. Deuxiéme liste de nombres chromosomiques nouveaux chez diverses
Dicotylédones et Monocotylédones d’ Afrique occidentale. Bull. Jard. Bot. (Bruxelles) 28:315—329.

. 1962. Enquéte sur les nombres chromosomiques dans une collection d’espéces tropicales. Rev. Cytol.
Biol. Vég. 25:411-447.

MANNING, J. C. AND F. M. GETLIFFE NorRIS. 1995.4. Duvernoia. In Flora of southern Africa, O. A. Leistner, ed.,
30(3, 1):15—17.

McDaAbDE, L. A. AND M. L. Moopy. 1999. Phylogenetic relationships among Acanthaceae: evidence from
noncoding trnL-trnF chloroplast DNA sequences. Amer. J. Bot. 86:70—80.

McDaAbDE, L. A., T. F. DANIEL, S. E. MASTA, AND K. M. RILEY. In press. Phylogenetic relationships within the
tribe Justicieae (Acanthaceae): evidence from molecular sequences, morphology, and cytology. Ann. Mis-
souri Bot. Gard.

PIOVANO, M. A. AND L. M. BERNARDELLO. 1991. Chromosome numbers in Argentinean Acanthaceae. Syst. Bot.
16:89-97.

PODLECH, D. 1986. Chromosomenstudien an Pflanzen des Saharo-sindischen Trockengebietes. Mitt. Bot.
Staatssamml. Miinchen 22:5—20.

RENARD, R., J. LAMBINON, M. REEKMANS, P. VAN DER VEKEN, AND M. GOVAERT. 1983. Nombres
chromosomiques de quelques angiospermes du Rwanda, du Burundi et du Kenya. Bull. Jard. Bot. Belg.
53:343-371.

SCOTLAND, R. W., J. A. SWEERE, P. A. REEVES, AND R. G. OLMSTEAD. 1995. Higher-level systematics of
Acanthaceae determined by chloroplast DNA sequences. Amer. J. Bot. 82:266—275.

VALSALA DEVI, G. AND P. M. MATHEW. 1982. /n IOPB chromosome number reports LX XVII. Taxon 31:773.

WELMAN, W. G. (compiler). 1993. Acanthaceae. /n Plants of southern Africa: names and distribution, T. H. Ar-
nold and B. C. de Wet, eds. Memoirs of the Botanical Survey of South Africa 62:652-665.

DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS jis)

APPENDIX

Species occurring in southern A frica for which chromosome numbers have been published previ-
ously. The taxon name is followed by the author of the name, the chromosome number(s) reported
(listed as n numbers), literature citation(s), and the country from which the plants studied came. The
provenance of the plants is based on information provided in the original publications. Two references
with chromosome numbers (both Ph.D. theses) have not been seen and the probable provenance is in-
dicated with a question mark and “not seen” follows. In other publications, where the source of mate-
rials studied is unclear or not provided, a question mark is indicated. The provenance of cultivated
plants noted below is unknown.

Acanthus pubescens (Oliv.) Engl., m = 28, Renard et al. 1983 (Burundi).

Asystasia gangetica (L.) T. Anderson (including 4. coromandeliana Nees), n = 13: Mangenot and Mangenot
1957 (western Africa), Mangenot and Mangenot 1962 (western Africa), Gadella 1977 (Cameroun), Ugborogho
and Adetula 1988 (Nigeria); 14: Subramanian and Govindarajan 1980 (India), Govindarajan and Subramanian
1983 (India); 22: Narayanan 1951a (India?); 24: Narayanan 1951a (India?); 25: De 1966 (India?), Sarkar et al.
1978 (India, cultivated); 26: Narayanan 195 1b (India), Grant 1955 (U.S. A., cultivated), Ellis 1962 (India), Kaur
1965 (India, cultivated), Valsala Devi and Mathew 1982 (India), Saggoo and Bir 1983 (India), Saggoo and Bir
1986 (India), Daniel 2000 (U.S. A., naturalized).

Barleria repens Nees, n = 20: Daniel and Chuang 1998 (cultivated).

Blepharis integrifolia (L.f.) E. Mey. (including B. molluginifolia Pers. and B. repens (Vahl) Roth), n = 17:
Kaur 1966 (India, probably cultivated), Sareen and Sanjogta 1976 (India), Ranganath 1981 (India? not seen),
Krishnappa and Ranganath 1982 (India), Ranganath and Krishnappa 1982 (India); n = 18: Govindarajan and
Subramanian 1983 (India), Subramanian and Govindarajan 1980 (India).

B. maderaspatensis (L.f.) Heyne ex Roth (including B. boerhaaviaefolia Pers.), n= 8: Valsala Devi and
Mathew 1982 (India), Saggoo 1983 (India?, not seen), Saggoo and Bir 1982a (India), 1983 (India), 1986 (India),
Govindarajan and Subramanian 1983 (India); n= 11: Ranganath 1981 (India? not seen), Krishnappa and
Ranganath 1982 (India), Ranganath and Krishnappa 1982 (India); n = 12: Ranganath and Krishnappa 1982 (In-
dia); n=13: Kaur 1966 (India), Sareen and Sanjogta 1976 (India); m=14: Vasudevan 1976 (India),
Subramanian and Govindarajan 1980 (India); n = 15: Miége 1962 (Senegal), Bir and Saggoo 1979 (India),
Saggoo 1983 (India? not seen), Bir and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1983 (India);
2n = 23: Ranganath and Krishnappa 1982 (India).

Crabbea velutina §. Moore (including C. reticulata C. B. Clarke), n = ca. 14: Renard et al. 1983 (Rwanda);
n= 21, Grant 1964 (cultivated).

Dyschoriste depressa Nees, n = 15: Govindarajan and Subramanian 1985 (India); n = 30: Bir and Saggoo
1979 (India), 1981 (India), Saggoo 1983 (India? not seen), Saggoo and Bir 1982b (India).

Elytraria acaulis (L.f.) Lindau, n = 17: Kaur 1969 (India?); 1 = 22: Govindarajan and Subramanian 1983 (In-
dia); n = 23: Subramanian and Govindarajan 1980 (India).

Hygrophila auriculata (Schum.) Heine (including Asteracantha longifolia), n= 16, De 1966 (India?),
Trivedi and Trivedi 1992 (India), Subramanyam and Kamble 1971 (India), Saggoo 1983 (India? not seen), Bir
and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1986 (India).

Hypoestes aristata R.Br., n = 30, Daniel and Chuang 1998 (cultivated).

H. forskaollii Vahl, n = 15, Podlech 1986 (Yemen).

156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

H. triflora Roem. and Schult., n = 15: Saggoo 1983 (India? not seen), Sagoo and Bir 1982a (India), 1983 (In-
dia).

Justicia anagalloides (Nees) T. Anderson, n= 9: Ensermu Kelbessa 1990 (Ethiopia); n= 18: Ensermu
Kelbessa 1990 (Ethiopia).

J. betonica L.,n = 14: Subramanian and Govindarajan 1980 (India), Narayanan 1951b (India?); n = 17: Ellis
1962 (India), Ranganath 1981 (India?, not seen), Bir and Saggoo 1979 (India), Krishnappa and Ranganath 1982
(India), Saggoo 1983 (India?, not seen), Valsala Devi and Mathew 1982 (India), Govindarajan and Subramanian
1983 (India), Bir and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1986 (India), Daniel and Chuang 1998
(cultivated), Daniel 2000 (U.S. A., naturalized).

J. exigua S. Moore, n = 7: Renard et al. 1983 (Rwanda), Ensermu Kelbessa 1990 (Ethiopia, Tanzania).

J. flava (Forssk.) Vahl, n = 13: Mangenot and Mangenot 1957 (western Africa), 1962 (western Africa),
Podlech 1986 (Yemen).

J. glabra Koenig ex Roxb., n = 27: Ranganath 1981 (India? not seen), Krishnappa and Ranganath 1982 (In-
dia).

J. matammensis (Schweinf.) Oliv., n = 7: Ensermu Kelbessa 1990 (Ethiopia).
J. odora (Forssk.) Lam., n = ca. 13: Hedrén 1989 (Tanzania).
Mackaya bella Harv., n = 42: Daniel and Chuang 1989 (cultivated).

Nelsonia canescens (Lam.) Spreng., n = 17: Saggoo 1983 (India? not seen), Saggoo and Bir 1983 (India),
n= 18: Daniel and Chuang 1993 (Panama).

Peristrophe paniculata (Forssk.) Brummitt (including P. bicalyculata (Retz.) Nees), n= 10: Narayanan
1951b (India); n= 15: Ahuja 1955 (India), Miége and Josserand 1972 (Senegal), Vasudevan 1976 (India),
Saggoo and Bir 1983 (India), 1986 (India), Verma and Dhillon 1967 (India).

Phaulopsis imbricata (Forssk.) Sweet, n = 16: Manktelow 1996 (Ethiopia, Tanzania, Madagascar); n = 17:
Daniel and Chuang 1998 (cultivated); n = ca. 32: Manktelow 1996 (Madagascar); 2n = 65: Manktelow 1996
(Malawi).

Rhinacanthus gracilis Klotz., n = 15: Daniel and Chuang 1998 (cultivated).

Ruellia cordata Thunb., n = 16, Rao and Mwasumbi 1981 (Tanzania, probably cultivated).

R. patula Jacq., n = 16: Baquar et al. 1966 (Pakistan), Rao and Mwasumbi 1981 (Tanzania, probably culti-
vated), Govindarajan and Subramanian 1983 (India), Subramanian and Govindarajan 1980 (India).

Ruspolia seticalyx (C. B. Clarke) Milne-Redh., n = 21: Daniel and Chuang 1998 (cultivated).

Thunbergia alata Boj.,n = 9: Grant 1955 (cultivated), Takizawa 1957 (cultivated), Shibata 1962 (Colombia,
naturalized), Saggoo 1983 (India?, not seen), Bir and Saggoo 1981 (India), Saggoo and Bir 1982b (India), 1986
(India), Daniel and Chuang 1989 (cultivated), Sugiura 1931 (cultivated?), Tijo 1948 (Indonesia), Narayanan
1951b (India), Kaur 1970 (India?); n = 16: Snoad 1952 (cultivated).

DANIEL, BALKWILL, AND BALKWILL: ACANTHACEOUS CHROMOSOME COUNTS 157

REFERENCES FOR APPENDIX

AHUJA, M. R. 1955. Chromosome numbers of some plants. Indian J. Genet. Pl. Breed. 15:142—-143.

BAQUAR, S. R., A HUSAIN, AND S. AKHTAR. 1966. Meiotic chromosome numbers in some vascular plants of
Indus Delta II. Bot. Notis. 119:24—32.

Bir, S. S. AND M. I. S. SAGGOO. 1979. Jn IOPB chromosome number reports LXV. Taxon 28:630-631.

. 1981. Cytopalynology of certain Acanthaceae and Labiatae. J. Palynol. 17:93—102.

DANIEL, T. F. 2000. Additional chromosome numbers of American Acanthaceae. Syst. Bot. 25:15—25.

DANIEL, T. F. AND T. 1. CHUANG. 1989. Chromosome numbers of some cultivated Acanthaceae. Baileya
23:86-93.

. 1993. Chromosome numbers of New World Acanthaceae. Syst. Bot. 18:283-289.

. 1998. Chromosome numbers of cultivated Acanthaceae and systematic implications. Pp. 309-330 in P.
Mathew and M. Sivadasan (eds.), Diversity and taxonomy of tropical flowering plants. Mentor Books,
Calicut.

DE, A. 1966. Cytological, anatomical and palynological studies as an aid in tracing affinity and phylogeny in the
family Acanthaceae. I. Cytological Studies. Trans. Bose Res. Inst. 29:139-175.

ELLIS, J. L. 1962. Chromosome numbers in some members of Acanthaceae. Sci. and Cult. 28:191—192.

ENSERMU KELBESSA. 1990. Justicia sect. Ansellia_(Acanthaceae). Acta Universitatis Upsaliensis Symbolae
Botanicae Upsalienses 29(2):1—96.

GADELLA, T. W. J. 1977. Jn IOPB chromosome number reports LVI. Taxon 26:259-261.

GOVINDARAJAN, T. AND D. SUBRAMANIAN. 1983. Karyomorphological studies in South Indian Acanthaceae.
Cytologia 48:491—504.

. 1985. Karyomorphological studies in South Indian Acanthaceae. Cytologia 50:473-482.

GRANT, W. F. 1955. A cytogenetic study in the Acanthaceae. Brittonia 8:121—149.

. 1964. In IOPB chromosome number reports. I. Taxon 13:108.

HEDREN, M. 1989. Justicia sect. Harnieria (Acanthaceae) in tropical Africa. Acta Universitatis Upsaliensis
Symbolae Botanicae Upsalienses 29(1):1—141.

KAvR, J. 1965. Chromosome numbers in Acanthaceae—II. Sci. and Cult. 31:531—532.

. 1966. Chromosome numbers in Acanthaceae 111 [sic]. Sci. and Cult. 32:142—143.

. 1969. Chromosome numbers in Acanthaceae—IV. Sci. and Cult. 35:61-63.

. 1970. Chromosome numbers in Acanthaceae—V. Sci. and Cult. 36:103—106.

KRISHNAPPA, D.G. AND R. M. RANGANATH. 1982. Jn IOPB Chromosome number reports LXXV. Taxon
31:364-365.

MANGENOT, S. AND G. MANGENOT. 1957. Nombres chromosomiques nouveaux chez diverses dicotylédones et
monocotylédones d’Afrique occidentale. Bull. Jard. Bot. (Bruxelles) 27:639-654.

. 1962. Enquéte sur les nombres chromosomiques dans une collection d’espéces tropicales. Rev. Cytol.
Biol. Vég. 25:41 1-447.

MANKTELOwW, M. 1996. Phaulopsis (Acanthaceae)—a monograph. Symb. Bot. Upsal. 31(2):1—184.

MIEGE, J. 1962. Quatriéme liste de nombres chromosomiques d’espéces d’ Afrique occidentale. Rev. Cytol. Biol.
Vég. 24:149-164.

MIEGE, J. AND N. JOSSERAND. 1972. Nombres chromosomiques d’espéces africaines et malgaches. Candollea
27:283-292.

NARAYANAN, C. R. 1951a. Nucleolar behaviour and chromosomal aberrations in mitosis of Acanthus ilicifolius
and Asystasia coromandeliana. Indian J. Genetics Pl. Breed. 11:205—210.

. 1951b. Somatic chromosomes in the Acanthaceae. J. Madras Univ. Bot. 21:220-231.

PODLECH, D. 1986. Chromosomenstudien an Pflanzen des Saharo-sindischen Trockengebietes. Mitt. Bot.
Staatssamml. Miinchen 22:5—20.

RANGANATH, R. M. 1981. Morphological and cytological studies in Acanthaceae. Ph.D. Thesis, Bangalore,
1981. [not seen]

RANGANATH, R. M. AND D.G KRISHNAPPA. 1982. Telocentric chromosomes and karyotypes in the genus
Blepharis. Caryologia 35:237—246.

RAO, P.N. AND L.B. MWASUMBI. 1981. /n IOPB chromosome number reports LX X. Taxon 30:79-80.

158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 12

RENARD, R., J. LAMBINON, M. REEKMANS, P. VAN DER VEKEN, AND M. GOVAERT. 1983. Nombres
chromosomiques de quelques angiospermes du Rwanda, du Burundi et du Kenya. Bull. Jard. Bot. Belg.
53:343-371.

SAGGOO, M. I. S. 1983. Cytomorphological studies on plants of economic importance of Bicarpellatae from In-
dia. Ph.D. Thesis, Punjabi Univ., Patiala. [not seen]

SAGGOO, M. I. S. AND S. S. BIR. 1982a. Jn IOPB chromosome number reports LX XVI. Taxon 31:574-598.

. 1982b. Cytological studies on certain Acanthaceae from central India. Proc. Indian Acad. Sci. (PI. Sci.)

91:479-486.

. 1983. Cytopalynological studies on Indian members of Acanthaceae and Labiatae. J. Palyn.

19:243-277.

. 1986. Meiotic studies in certain members of family Acanthaceae from South India. J. Indian Bot. Soc.
65:310-315.

SAREEN, T. S. AND K. SANJOGTA. 1976. Chromosome numbers in some species of Acanthaceae. Cytologia
41:283-290.

SARKAR, A.K., M. CHAKRAVERTY, N.C. SAHA, S. K. DAS, AND D. HAZRA. 1978. Jn IOPB chromosome number
reports LXII. Taxon 27:519-535.

SHIBATA, K. 1962. Estudios citologicos de plantas colombianas silvestres y cultivadas. J. Agricul. Sci. 8:49-62.

SNOAD, B. 1952. Chromosome counts of species and varieties of garden plants. Annual Report, John Innes Horti-
cultural Institution 42:47—S0.

SUBRAMANIAN, D. AND T. GOVINDARAJAN. 1980. Cytotaxonomy of some species of Acanthaceae. J: Cytol.
Genet. 15:90—92.

SUBRAMANYAM, K. AND N.P. KAMBLE. 1971. /n IOPB chromosome number reports XX XI. Taxon 20:160.

SuGIURA, T. 1931. A list of chromosome numbers in angiospermous plants. Bot. Mag. (Tokyo) 45:353-355.

TAKIZAWA, Y. 1957. Die Struktur der Pachytanchromosomen einiger Acanthaceen, sowie eine Reihe neu
bestimmter Chromosomenzahlen. Cytologia 22:118—126.

Tuo, J. H. 1948. The somatic chromosomes of some tropical plants. Hereditas 34:135—146.

TRIVEDI, M. P. AND R. N. TRIVEDI. 1992. Chromosomal behaviour in weeds. Proc. Conf. Cytol. and Genet.
3:188-198.

UGBOROGHO, R. E. AND O. A. ADETULA. 1988. The biology of the Asystasia gangetica complex (Acanthaceae)
in Lagos State, Nigeria. Feddes Rep. 99:507-517.

VALSALA DEVI, G. AND P.M. MATHEW. 1982. /n IOPB chromosome number reports LX XVII. Taxon 31:773.

VASUDEVAN, K. N. 1976. Contribution to the cytotaxonomy and cytogeography of the flora of the western Hima-
layas (with an attempt to compare it with the flora of the Alps). Part III. Ber. Schweiz. Bot. Ges. 86:152—203.

VERMA, S. C. AND S. S. DHILLON. 1967. /n IOPB chromosome number reports XI. Taxon 16:221—222.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 13, pp. 159-169, 8 figs. f ene. December 7, 2000

A New Species of the Soft Coral Genus Eleutherobia Putter, 1900
(Coelenterata: Alcyoniidae) from the Tonga Islands

by

Gary C. Williams
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

The soft coral genus Eleutherobia (family Alcyoniidae) was previously known to contain
fifteen valid species, and is distributed from eastern and southern Africa to the western Pa-
cific. A new species is here described from Tonga, thereby making a total of sixteen species,
and extending the known range of the genus approximately 3000 km to the southeast into
the central South Pacific. The new taxon is superficially similar to Eleutherobia grayi(J. A.
Thomson and Dean, 1931), from the Indonesian and Ryukyuan archipelagos, but is distin-
guished from it and all other species by a unique complement of sclerite types.

The Indo-Pacific and southern African soft coral genus Eleutherobia is noteworthy for recently
discovered, bioactive, natural products that have been isolated from two Indian Ocean species. These
compounds include novel diterpenes (Hooper et al. 1997) and the cytotoxic agent eleutherobin (Long
et al. 1998). The latter compound has recently been found to have anticancer properties and has re-
cently been synthesized (Xiao et al. 1999). The diterpenes are produced by Eleutherobia lutea
Benayahu and Schleyer, 1995, from the east coast of South Africa, and eleutherobin comes from an
undescribed species off the west coast of Australia.

Of the fifteen described species that are considered valid for the genus Eleutherobia, twelve in-
habit the Indo-West Pacific from East Africa to Japan and Indonesia, and three are considered en-
demic to southern Africa [ Verseveldt and Bayer (1988) and the present paper]. Verseveldt and Bayer
(1988) provided a taxonomic revision of the genus Eleutherobia, Williams (1992a) reviewed the
southern A frican species, and Benayahu and Schleyer (1995) described a new species from South Af-
rica. Color photographs of living soft corals in the genus are provided by Williams (1996:34) and
Benayahu and Schleyer (1995:2).

A recently discovered species from Tonga is here described. It differs from other members of the
genus by having a sclerite complement that includes very slender capstans with thin radiating whorls
of tubercles, a relatively large number of crosses > 0.10 mm in length (many of which have elongated
tapering points), and a scarcity of triradiate forms or needle-like spindles. The addition of the new spe-
cies makes a total of sixteen species of Eleutherobia recognized as valid—three from southern Africa
and thirteen from the Indo-West Pacific.

METHODS

Material was collected by SCUBA and preserved in 70% ethanol. Sclerites were isolated using
sodium hypochlorite. Micrographs were made using a Kodak MDS100 digital video camera and a

159

FEB 95 2001

160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 13

Wild M400 photomicroscope. Scanning electron micrographs were made on Hitachi S-510 and Leo
1400 Series scanning electron microscopes. Abbreviations used in the text are as follows: CAS (Cali-
fornia Academy of Sciences, San Francisco), CRRF (Coral Reef Research Foundation, Palau).

SYSTEMATIC ACCOUNT

Family Alcyontidae Lamouroux, 1812
Eleutherobia Pitter, 1900

Eleutherobia Pitter, 1900:449. Verseveldt and Bayer, 1988:27. Williams, 1992:306.
Nidalia (in part): non Gray, 1835.

Bellonella (in part): non Gray, 1862.

Metalcyonium (in part): non Pfeffer, 1889.

DIAGNOSIS. — Alcyoniid soft corals with colonies digitiform, finger-shaped, often conical and
tapering, rarely multilobate. Polyps monomorphic. Calyces absent, although retracted polyps often
form low rounded to conspicuous protuberances. Sclerites mostly derived from radiates, although
spindles, crosses, barrels, or tuberculate spheroids may also be present.

TYPE SPECIES. — Eleutherobia japonica Pitter, 1900, by monotypy.

DIVERSITY AND DISTRIBUTION. — Sixteen species of the Indo-West Pacific (southern and east-
ern Africa to Japan and Tonga). |

Eleutherobia zanahoria sp. nov.
Figs. 1-8

MATERIAL EXAMINED. — HOLOTYPE: CAS 118501, station number CRRF #OCDN 5464-X,
Tonga, southwest of Vavau, south side of Kitu channel, a small channel between Kitu Island and north
side of Nuapapu Island (18° 41.25’S, 174° 04.05'W), 25-30 m depth, 12 November 1997, collected
by Coral Reef Research Foundation, one whole specimen, 64 mm in length. PARATYPES: CAS
118502, same data as holotype, one whole specimen, 63 mm in length. CAS 118503, same data as
holotype, one specimen cut in half longitudinally, 53 mm in length.

DIAGNOSIS. — Alcyoniid soft corals with digitiform to lobate colony shape. Several finger-like
lobes may be united by a common basal holdfast. Stalk very short, polyps distributed over approxi-
mately 95% of each colony. Sclerites are radiates, crosses, and irregular forms presumably derived
from radiates; some crosses with finely attenuated tips. Polypary sclerites relatively gracile,
0.04—0.12 mm long; holdfast and stalk sclerites more robust, 0.07—0.16 mm long. Polyp sclerites ab-
sent. External coenenchymal color carrot orange throughout. Sclerites pale yellow-orange.

DESCRIPTION. — Growth form and size: The wet-preserved holotype is unbranched and fin-
ger-shaped, elongate conical, gradually tapering from proximal base to distal end. The apex is gently
rounded. The basal holdfast has the largest width (20 mm), while the apex region has the smallest
width (5 mm). The polypary comprises over 95% of the total colony length, as the polyps begin to ap-
pear immediately above the holdfast (Fig. 1). Wet-preserved paratype CAS 118702 is 63 mm in
length, and lobate with four terminal lobes arising from two trunks that are unified into a single trunk
and holdfast at the base (Fig. 2). The widest portion of the specimen is in the holdfast region (31 mm),
while the narrowest lobe is 4 mm in width just below its apex. The lobes vary from 8 to 28 mm in
length. They are mostly cylindrical in shape and slightly curved with gently rounded ends.
Wet-preserved paratype CAS 118503 is 53 mm in length and similar in shape to the holotype. The
widest portion at the base measures 9 mm, while the narrowest portion near the apex is 3 mm in width

WILLIAMS: NEW SPECIES OF SOFT CORAL

FIGURE 1. Eleutherobia zanahoria sp. nov. A. Underwater photograph of living soft corals at the type locality. Photograph by
Pat Colin, courtesy of the Coral Reef Research Foundation, Palau. B. Wet-preserved holotype (CAS 118501), 64 mm in length.
C. Wet-preserved paratype (CAS 118503), 53 mm in length; cut longitudinally into two halves; scale bar = 16 mm.

162 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 13

(Fig. 1C). In both paratypes, as in the
holotype, polyps appear immediately above
the holdfast region, so that it is difficult to
distinguish a conspicuous stalk region.

Polyps. The polyps are retracted into
the colonies in all specimens and are thus
not observable. A piece of surface co-
enenchyme was removed in paratype CAS
118503 to show the arrangement of several
retracted polyps (Fig. 3). A single polyp that
was isolated from paratype CAS 118503 and
dissolved in sodium hypochlorite, revealed
no sclerites in the anthocodial region or
polyp walls. Calyces are absent, the re-
tracted polyps form recessed slits or depres-
sions on the surface of the polypary, or they
are flush with the surrounding polypary sur-
face.

Sclerites. The sclerites are densely set
in the surface and subsurface coenenchyme
of the polypary and holdfast region, as well
as in the deep interior of the holdfast region.
Sclerites are absent from the polyps and
walls of the gastric cavities. The sclerites are
radiates, crosses, and irregularly-shaped
forms that are presumably derived from ra-
diates, 0.04—0.17 mm in length. The scler-
ites of the polypary are more gracile and, for
the most part, somewhat smaller than
sclerites of the stalk and holdfast region.
Sclerites from the polypary surface are
mostly slender radiates, 0.05—0.09 mm in

length (Figs. 4A, 5). Sclerites from the

- eens ea ig ae ia sp. nov. Paratype (CAS subsurface of the polypary are similar radi-

ates, including some crosses and irregular

forms, 0.04—0.12 mm in length (Figs. 5,

6B). Sclerites from the surface of the holdfast region are relatively robust radiates, crosses, and irregu-

lar forms, 0.07—0.17 mm long (Figs. 4B, 7). Sclerites from the deep interior of the holdfast region are

robust, mostly irregular forms derived from and similar to radiates, with some radiates and crosses,
0.08—-0.16 mm long (Figs. 6A, 7).

Color. The coenenchyme is vivid orange throughout (Fig. 1); polyps are creamy white. Sclerites
are pale orange or yellowish, some are colorless.

ETYMOLOGY. — The specific epithet is derived from the Spanish, zanahoria (a carrot); in refer-
ence to the carrot-like appearance and color of this soft coral.

DISTRIBUTION. — Known from the type locality—the Tonga Islands in the central, South Pacific
Ocean (Fig. 8). This species is also reported to occur on the Great Barrier Reef, Queensland, Australia
(P. Alderslade, pers. comm.).

WILLIAMS: NEW SPECIES OF SOFT CORAL 163

FiGurE 3. Eleutherobia zanahoria sp. nov. Micrograph of paratype (CAS 118503) with portion of surface coenenchyme re-
moved, showing interior of colony and several retracted polyps. Abbreviations: gc - gastric cavity; ic - interior coenenchyme; rp
- retracted polyp; sc - surface coenenchyme; t - tentacle. Scale bar = 1.3 mm.

DISCUSSION

Species of the genus Eleutherobia closely resemble those of another Indo-West Pacific genus,
Paraminabea Williams and Alderslade, 1999. The two taxa differ however, in that species of
Eleutherobia are monomorphic, while those of Paraminabea have dimorphic polyps (Williams
1992b:5, fig. 2C; Williams and Alderslade 1999:347, fig. 6B). Unfortunately, siphonozooids are of-
ten very difficult to observe in preserved material. This circumstance has been at least partly responsi-
ble for the misidentification of soft coral material in the past, and will no doubt result in similar
mistakes being made in the future. Thin sectioning of the surface and subsurface coenenchyme of the
polypary of the soft coral in question, together with the microscopic examination of these sections, is
often necessary to detect siphonozooids in tightly contracted and retracted preserved material.

Of the twenty valid species of the genus Eleutherobia, E. zanahoria sp. nov. is superficially most
similar to Eleutherobia grayi (J. A. Thomson and Dean 1931), redescribed by Verseveldt and Bayer
(1988:33), from Indonesia and the Ryukyu Islands. The two species can be differentiated as follows.
In wet-preserved specimens of E. grayi, the retracted polyps often form low rounded, to more pro-
nounced and mammiform, to conical protuberances on the surface of the polypary. In E. zanahoria sp.
nov., on the other hand, retracted polyps in preserved specimens or retracted living animals, appear as
shallow slits or pits, or are flush with the surface of the polypary (Fig. 1). Although sclerite size is simi-

164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 13

FiGure 4. Eleutherobia zanahoria sp. nov. A. Sclerites from the surface of the polypary. B. Sclerites from the surface of the
holdfast region of the stalk. Scale bars = 0.1 mm.

lar in both species, the complement of sclerites differs. In E. grayi, triradiates are relatively common
and the crosses (quadriradiates) that are found, are relatively small (< 0.06 mm long), and have short
and knobby or truncated rays (Verseveldt and Bayer 1988:93, fig. 24). By comparison, in E.
zanahoria sp. nov.., triradiates are rare or absent, and many crosses are relatively large (up to 0.14 mm
in length), and have finely tapered and acute-tipped rays (Figs. 4B, SA, 6A, 7A—B, 7F). Finally, in re-
gards to coloration, the two species differ in that E. grayi is bicolored (red or red-orange with yellow
retracted polyp mounds, while E. zanahoria sp. nov. is uniform bright orange throughout (Fig. 1).

WILLIAMS: NEW SPECIES OF SOFT CORAL 165

FiGureE 5. Eleutherobia zanahoria sp. nov. Scanning electron micrographs of sclerites from the polyparium. A. 0.09 mm; B.
0.05 mm; C. 0.07 mm; D. 0.05 mm; E. 0.07 mm; F. 0.06 mm; G. 0.06 mm; H. 0.06 mm; I. 0.06 mm; J. 0.06 mm; K.0.09 mm.

166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 13

Ficure 6. Eleutherobia zanahoria sp. nov. A. Sclerites from the interior of the holdfast. B. Sclerites from the interior of the
polypary. Scale bar for A and B = 0.1 mm.

ACKNOWLEDGMENTS

Iam grateful to Alan Leviton and Katie Martin (Scientific Publications, California Academy of
Sciences), Leen van Ofwegen (National Museum of Natural History, Leiden), and Phil Alderslade
(Northern Territory Museum of Arts and Sciences, Darwin) for their comments and suggestions. I
also thank Delisse Ortiz and Angel Valdés (Department of Invertebrate Zoology and Geology, Cali-
fornia Academy of Sciences) for derivation of the specific epithet.

Field work was made possible by a contract from the United States National Cancer Institute to
Pat and Lori Colin, Coral Reef Research Foundation, Palau.

WILLIAMS: NEW SPECIES OF SOFT CORAL

FIGURE 7. Eleutherobia zanahoria sp. nov. Scanning electron micrographs of sclerites from the holdfast region of the stalk.
A. 0.14 mm; B. 0.10 mm; C. 0.09 mm; D. 0.08 mm; E. 0.08 mm; F. 0.10 mm; G. 0.08 mm; H. 0.08 mm; I. 0.10 mm; J.
0.08 mm; K. 0.10 mm; L. 0.09 mm.

168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 13

FIGURE 8. Map of the Indo- West Pacific showing geographic distribution of the genus Eleutherobia. Arrow shows type local-
ity of Eleutherobia zanahoria sp. nov.

LITERATURE CITED

BENAYAHU, Y. AND M. H. SCHLEYER. 1995. Corals of the south-west Indian Ocean II. Eleutherobia aurea spec.
nov. (Cnidaria, Alcyonaria) from deep reefs on the KwaZulu-Natal Coast, South Africa. South African As-
sociation for Marine Biological Research, Oceanographic Research Institute, Investigational Report
68:1-12.

GRAY, J. E. 1835. Characters of a new genus of corals (Nidalia). Proceedings of the Zoological Society of Lon-
don 3:59-60.

. 1862. Description of two new genera of zoophytes (Solenocaulon and Bellonella) discovered on the
north coast of Australia by Mr. Rayner. Proceedings of the Zoological Society of London 1862:34-37.

Hooper, G. J., M. T. DAVIES-COLEMAN, AND M. SCHLEYER. 1997. New diterpenes from the South African soft
coral Eleutherobia aurea. Journal of Natural Products 60:889-893.

LONG, B. H., J. M. CARBONI, A. J. WASSERMAN, L. A. CORNELL, A. M. CASAZZA, P. R. JENSEN, T. LINDEL, W.
FENICAL, AND C. R. FAIRCHILD. 1998. Eleutherobin, a novel cytotoxic agent that induces tubulin polymer-
ization, is similar to paclitaxel (Taxol). Cancer Research 58:1111—1115.

PFEFFER, G. 1889. Zur Fauna von Sud-Georgien. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten
6(2):49-SS.

PUTTER, A. 1900. Alcyonaceen des Breslauer Museum. Zoologische Jahrbiicher (Systematik) 13(5):443-462.

THOMSON, J. A. AND L. M. I. DEAN. 1931. The Alcyonacea of the Siboga Expedition with an addendum to the
Gorgonacea. Siboga Expedition Monographs 13d:1—227.

VERSEVELDT, J. AND F.M. BAYER. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and
Nidaliopsis (Octocorallia: Alcyoniidae and Nidalliidae), with descriptions of two new genera. Zoologische
Verhandelingen 245:1—131.

WILLIAMS: NEW SPECIES OF SOFT CORAL 169

WILLIAMS, G.C. 1992a. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals
(Coelenterata, Anthozoa). Annals of the South African Museum 100(3):249-358.

. 1992b. Revision of the soft coral genus Minabea (Octocorallia: Alcyoniidae) with new taxa from the

Indo-West Pacific. Proceedings of the California Academy of Sciences 48(1): 1-26.

. 1996. Octocorallia—Octocorals. Pp. 32-60 in Coral reef animals of the Indo-Pacific—animal life from
Africa to Hawai’i exclusive of the vertebrates. T. M.Gosliner, D. W. Behrens, and G. C. Williams. Sea
Challengers, Monterey. 314 pp .

WILLIAMS, G. C. AND P. ALDERSLADE. 1999. Revisionary systematics of the western Pacific soft coral genus
Minabea (Octocorallia: Alcyoniidae), with descriptions of a related new genus and species from the
Indo-Pacific. Proceedings of the California Academy of Sciences 51(7):337-364.

XIAO, T. C., S. K. BHATTACHARYA, B. ZHOU, C. E. GUTTERIDGE, T. R. R. PETTUS, AND S. J. DANISHEFSKY.
1999. The total synthesis of eleutherobin. Journal of the American Chemical Society 121(28):6563-6579.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE

LIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 14, pp. 171-181, 5 figs. | December 7, 2000

DEC 11 2000

ee |
Two New Species of Aldisa eral!" (Mollusea, Nudibranchia)

from the Tropical Indo-Pacific

by

Hillary R. Elwood, Angel Valdés and Terrence M. Gosliner
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

This paper provides descriptions of two new species of the genus Aldisa from the tropical
Indo-Pacific. Aldisa albatrossae sp. nov. from Japan, the Philippines, and Indonesia, is
characterized by having a blue-green dorsum with distinctive black markings and blotches
of yellow-orange color. Aldisa williamsi sp. nov. from Papua New Guinea and Indonesia,
has a bluish dorsum with a circular black mark. The two species are distinguishable based
on differences in body coloration, arrangement of tubercles and characters of the repro-
ductive system. Other species that have similar color patterns and also appear to be mimics
of phylllidid nudibranchs are discussed. The new species appear to have two plesiomorphic
traits that were previously undescribed for the genus Aldisa, unipinnate gills (only in A.
williamsi), and presence of two large hamate radular teeth (in both species).

The genus A/disa is characterized by having elongate denticulate teeth, penial spines, conical tu-
bercles and absence of oral tentacles. The genus A/disa presently contains twelve valid species,
mainly found in the cold-temperate waters of the Atlantic Ocean. Eleven species were identified as
valid during the revision of this genus by Millen and Gosliner (1985) and one species has been de-
scribed since, Aldisa barlettai Ortea and Ballesteros, 1988 from the Cape Verde Islands. Only one of
these twelve previously described species is from the tropical Indo-Pacific, Aldisa pikokai Bertsch
and Johnson, 1967, from Hawaii.

The objective of this paper is to describe two new species from the tropical Indo-Pacific. The de-
scriptions are based on specimens deposited in the Department of Invertebrate Zoology of the Califor-
nia Academy of Sciences (CASIZ).

SPECIES DESCRIPTIONS

Aldisa albatrossae sp. nov.
Figs. 1A—B, 2, 3

Doris sp. 1 Ono, 1999:138, fig. 178.
Chromodoris sp. Masuda, 1999:195, bottom photo.

MATERIAL EXAMINED. — HOLOTYPE: Seragaki Beach, 1.3 km ENE of Maeki-zaki, Okinawa,
Ryukyu Islands, Japan (26°30.4’N, 127°52.6'E), 3 April 1993, 19 mm long, collected by Robert
Bolland (CASIZ 89034). PARATYPES: Seragaki Beach, 1.3 km ENE of Maeki-zaki, Okinawa,
Ryukyu Islands, Japan (26°30.4'N, 127°52.6’E), 11 April 1993, 1 specimen, 21 mm long, collected
by Robert Bolland (CASIZ 88884); Horseshoe Cliffs, 1 km WNW of Onna Village, Okinawa,

171

172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 14

Ryukyu Islands, Japan (26°30.0'N, 127°50.9’E), 9 May 1998, Ispecimen, 17 mm long, collected by
Robert Bolland (CASIZ 115720); Seragaki Tombs, Okinawa, Ryukyu Islands, Japan (26°30.4'N,
127°52.6’E), 29 April 1995, 2 specimens, 22 mm long and 19 mm long, collected by Robert Bolland
(CASIZ 105287); Horseshoe Cliffs, 1 km WNW of Onna Village, Okinawa, Ryukyu Islands, Japan
(26°30.0'N, 127°50.9’E), 26 March 1994, 1 specimen, 11 mm long, collected by Robert Bolland
(CASIZ 99086).

ETYMOLOGY. — Aldisa albatrossae gets its name from the Albatross expedition of 1907—10 to
the Philippines in which a specimen of this species was collected and illustrated (Fig. 1A).

GEOGRAPHIC RANGE. — So far this species is known from the Kerama and Ryukyu Islands of Ja-
pan, the Albatross specimen collected in the Philippines, and from a photograph of a species from Ko-
modo Island, Indonesia taken by Jim Black.

EXTERNAL MORPHOLOGY. — The living animals are 1 1—21 mm in length. The dorsum of the liv-
ing animal is a bluish color (Fig. 1B). This color is darker at the edges and lighter towards the middle
of the dorsum. It is interrupted by the presence of numerous white tubercles. The dorsum also contains
a distinctive black pattern, which is virtually identical in all specimens examined. The black pattern
begins as a T-shape crossing in front of and between the rhinophores. Behind the rhinophores, the
T-shape connects with a black rectangular mark that spans the dorsum to the branchial pocket. Two
black lines extend from the rectangular mark around the sides of the branchial pocket. In one specimen
the black marks form a continuous band around the posterior margin of the branchial sheath. These
characteristic black markings on the dorsum can also be seen in the preserved specimens. Yel-
low-orange splotches are present on the dorsum of the living animal. These marks of yellow-orange
lie at the front edges of the T of the black mark and along the anterior edge of the rectangular portion of
the black mark. Yellow-orange lines also extend from the sides of the branchial pocket to the posterior
edges of the dorsum.

The rhinophores are dark, uniformly off-white and lamellate, containing 18 lamellae ina 21 mm
long specimen. The branchial leaves are a darker gray than the rhinophores and range from 6—9 in
number. They are bipinnate and relatively sparsely branched. .

There are numerous rounded tubercles over the surface of the body. Four to five rows of tubercles
are found outside of the broad black ring on all sides of the body. An additional four to five rows are
found within the black band. The tubercles of the body are smaller towards the edges of the dorsum
(Fig. 2D) with a row of small tubercles around the entire margin. Around the branchial pocket, there
are two rows of tubercles. The tubercles nearest the gill are equally spaced and of alternating large
then small size. The other row also consists of tubercles of alternating sizes. Each rhinophore is sur-
rounded by four tubercles. These tubercles also alternate large and small with the larger tubercles on
the left and right sides of each rhinophore. All tubercles are opaque white and conical (Fig. 2D).

The anterior portion of the foot is grooved but not notched. The head has short oral protrusions
(Fig. 3C) with a short grove along their outer edge.

RADULA. — The radular formula is undeterminable owing to the elongate, overlapping teeth.
The radular teeth are narrow and elongate with broad triangular bases (Fig. 2A). The top third or fourth
of each tooth bears 20-24 sharp denticles, which extend along the side of each tooth. The denticles
near the apex of the teeth are much longer than those on the sides (Fig. 2B). There are also two large,
hamate teeth, each one situated on one half-row, which lack denticles (Fig. 2C).

REPRODUCTIVE SYSTEM. — The ampulla is tubular and thick, winding back on itself before nar-
rowing into a thin tube that connects with the oviduct and the prostate (Fig. 3A). The oviduct connects
to the female gland underneath the prostate. The prostate is granular, tubular and slightly larger than
the ampulla. Its end joins with the thin deferent duct. The deferent duct loops before meeting the vagi-
nal duct at the genital atrium. The genital atrium is no wider than the width of the ducts and extends
only slightly beyond their meeting. The penial bulb is armed with 13 rows of six hooks each. The
hooks have a wide base tapering off to a thin cusp (Fig. 3B). The vaginal duct is about the same length

ELWOOD, VALDES, AND GOSLINER: ALDISA 173

FiGure 1. Color illustrations of the living animals. A. Unpublished painting of Aldisa albatrossae from the Albatross Expe-
dition; B. Living paratype of A. albatrossae sp. nov. from the Ryukyu Islands, Japan (CASIZ 88884); C. Living paratype of A.
williamsi sp. nov. from Barracuda Point, Papua New Guinea (CASIZ 109791).

174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 14

and width as the deferent duct. The vaginal duct runs underneath the seminal receptacle to connect to
the round bursa copulatrix. Another duct connects the bursa copulatrix to the smaller, stalked seminal
receptacle. The short uterine duct connects to the middle of this duct and leads into the female gland.

Aldisa williamsi sp. nov.
Figs. 1C;.4, 5

Doris? sp. 10 Rudman, 2000.

MATERIAL EXAMINED. — HOLOTYPE: Barracuda Point, E side of Tab Island, near Madang, N
coast of Papua New Guinea, 14 November 1990, 11 mm preserved length, collected by Terry
Gosliner (CASIZ 075936). PARATYPE: Bomber Reef, Madang, Papua New Guinea, 2 November
1996, 1 specimen 14 mm preserved length, collected by Gary Williams (CASIZ 109791).

ETYMOLOGY. — This species is named after our good friend and colleague, Gary Williams
(CASIZ). He collected one of the specimens of this new species and a number of other nudibranchs
during several expeditions to the tropical Indo-Pacific.

GEOGRAPHIC RANGE. — This species has only been collected at Madang, Papua New Guinea,
but there are records from two photographs of specimens from Sulawesi, Indonesia taken by Lindsay
Warren (Rudman, 2000).

EXTERNAL MORPHOLOGY. — The dorsum of the living animal is a uniform bluish gray color
with opaque white tubercles (Fig. 1C). There is a distinctive black pattern on the dorsum of the living
animal that can also be seen in preserved specimens. This pattern consists of an oval line that stretches
from in front of the rhinophores to behind the branchial pocket. In the two specimens examined there
is also a short line that runs between the two rhinophores and connects posteriorly to the oval line be-
hind the left rhinophore.

The rhinophores are a pale brown color with a bluish base. They have 13 lamellae in a 14 mm
long preserved specimen. The gill consists of six unipinnate branchial leaves. The two posteriormost
leaves are bifurcated at the base. They are a light yellow hue with splotches of opaque white.

The tubercles of the dorsum are opaque white and are of roughly equal size. Smaller tubercles are
present at the edges of the dorsum. There are two to three rows of tubercles outside of the broad black
band that encircles the notum with an additional two to three rows inside the band. Around each
rhinophore, there are two tubercles. In the paratype specimen the outer tubercle is the same pale brown
color as the rhinophore. The inner tubercle is opaque white. In the holotype, both tubercles are opaque
white. Around the branchial pocket, there is a single row of 8 white tubercles of alternating large and
small size. These tubercles are evenly spaced between the branchial leaves and are different from
those of the dorsum.

The anterior portion of the foot is not notched. The head contains short oral protrusions (Fig. 5C)
and these protrusions have a groove along the outer edge.

RADULA. — The radular formula is undeterminable. The radular teeth are narrow and elongate
with broad triangular bases (Fig. 4A). The top third or fourth of each tooth bear 15—20 sharp denticles,
which extend along the side of each tooth. The denticles near the apex of the teeth are much longer
than those on the sides (Fig. 4B). There are also two large, hamate teeth, each one situated on one
half-row (Fig. 4C), which lack denticles.

REPRODUCTIVE SYSTEM. — The ampulla is tubular and convoluted (Fig. 5A). It narrows into a
thin tube and connects to the oviduct and prostate. The prostate is longer and slightly thinner than the
ampulla. It loops a few times before connecting to the thin deferent duct. The deferent duct and vaginal
duct connect at the genital atrium. The two ducts are of roughly equal width and length. The penial
bulb is armed with 11 rows of 8 penial hooks each. The hooks have a wide base and a relatively short
cusp (Fig. 5B). The vaginal duct connects to the large round bursa copulatrix. A separate duct con-

ELWOOD, VALDES, AND GOSLINER: ALDISA 175

FIGURE 2. Aldisa albatrossae sp. nov. (CASIZ 88884) scanning electron micrographs. A. Elongate radular teeth, scale
bar = 10 pm; B. Heads of elongate radular teeth, scale bar = 2 um; C. Hamate tooth, scale bar = 20 um; D. Tubercles of dorsum,
scale bar = 200 pm.

nects to the short stalk of the seminal receptacle. The very thin uterine tube connects to this duct and
then connects to the female gland near the genital atrium.

DISCUSSION

The revision of the genus A/disa by Millen and Gosliner (1985) produced a diagnosis of the genus
with the following characteristics: elongate radular teeth with multiple denticulations, a dorsum with
low conical tubercles (except for A. pikokai, which lacks dorsal tubercles), a ring of tubercles around
branchial and rhinophoral pockets, bipinnate or tripinnate branchial leaves, penial spines in almost all

176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 14

FiGuRE 3. Aldisa albatrossae sp. nov. (CASIZ 88884). A. Reproductive system, scale bar = 0.5 mm; B. Penial hooks, scale
bar = 5 pm; C. Ventral view of the anterior border of the foot, scale bar = 1 mm. Abbreviations: am, ampulla; be, bursa
copulatrix; dd, deferent duct; fg, female gland; pr, prostate; sr, seminal receptacle; v, vagina.

species, absence of oral tentacles and an anterior portion of the foot that is not notched. The two spe-
cies described in this paper exhibit almost all the characteristics of the genus A/disa noted above and
are identified as such.

They join the 12 previously described species of Aldisa. The species previously described in-
clude the 11 species identified as valid during revision of Aldisa by Millen and Gosliner (1985) and
one species that has been described since, Aldisa barlettai Ortea and Ballesteros, 1988, from the Cape
Verde Islands. While the species described in this paper are clearly placed within the genus Aldisa,
they differ markedly from the species previously described.

Aldisa albatrossae and A. williamsi both exhibit a character that, before now, has not been de-
scribed for the genus A/disa: the presence of a pair of unique teeth in their radula. As mentioned be-
fore, these two teeth are short, wide, smooth and hamate, differing from the numerous thin, elongate
teeth, characteristic of Aldisa, that make up the rest of the radula. The presence of these two broad

ELWOOD, VALDES, AND GOSLINER: ALDISA

FiGuRE 4. Aldisa williamsi, sp. nov. (CASIZ 109791) scanning electron micrographs. A. Elongate radular teeth, scale
bar = 10 pm; B. Heads of elongate radular teeth, scale bar = 1 pm; C. Hamate tooth, scale bar = 10 pm.

teeth in these species is believed to be a plesiomorphic state due to the resemblance of these teeth to the
hamate teeth of the other cryptobranch dorids. Also the unipinnate branchial leaves of A. williamsi,
which occur in some basal dorids, may well be a plesiomorphy within A/disa.

The coloration of A. albatrossae and A. williamsi also distinguishes them from the 12 species of
Aldisa previously described. None of the 11 species that Millen and Gosliner (1985) identify as valid

178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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exhibit any of the bright colors or black markings found on the notum of the new species. Aldisa
barlettai from the Cape Verde Islands is the only other species to exhibit a bright and multi-colored
body. However, the coloration of A. barlettai, which has a dorsum of blue-violet with white and or-
ange tubercles (Ortea and Ballesteros 1988), is quite different from the new species which have a
blue-green dorsum, white tubercles, black markings, and, in the case of A. albatrossae, yellow-orange
splotches. Both species were collected in the warm waters of the Indo-Pacific, A. albatrossae from
around the Kerama and Ryukyu Islands of Japan, the Philippines and Indonesia and A. williamsi from
Papua New Guinea and Indonesia. Aldisa pikokai, from Hawaii, is the only other species found in the
tropical Indo-Pacific. The orange-red coloration and absence of tubercles on the dorsum of A. pikokai
(Bertsch and Johnson 1982), readily distinguishes it from the new species.

Aldisa albatrossae and A. williamsi share some characteristics of the radula, reproductive system
and coloration, but there is little doubt that they are indeed distinct species.

Aldisa albatrossae is known from the six specimens examined in this study from Okinawa. Pho-
tographs of an additional three specimens from Okinawa were provided by Bob Bolland. Two addi-
tional photographs of specimens from the Kerama Islands appear in two Japanese books (Ono 1999;
Masuda 1999). A single specimen was depicted in the drawing of the Albatross specimen from the
Philippines (Fig. 1A). An additional specimen was photographed by Jim Black from Komodo Island,
Indonesia. In all of these 13 specimens there are consistent elements of the external morphology. A
black T-shaped marking is present on the anterior end of the notum and continues posteriorly between
the rhinophores. This band then bifurcates and continues posteriorly as two bands that reconnect in
front of the branchial sheath. In one specimen a second black band encircles the posterior portion of
the branchial sheath. In all specimens, anterior and posterior patches of yellow-orange are present. In
all of these specimens there are four tubercles surrounding each rhinophore sheath. In all specimens
there are two bands of tubercles surrounding along the margin and immediately ventral to the
branchial sheath. All specimens of A. albatrossae have bipinnate gill branches.

Aldisa williamsi is described here from two specimens from Papua New Guinea. Two additional
specimens from Sulawesi, Indonesia, depicted by Rudman (2000) as Doris? sp. 10, also appear to be
the conspecific with A. williamsi. These two specimens differ from the Papua New Guinea specimens
in two minor details. In the specimens from New Guinea only one black band encircles the left
rhinophore. In the specimens from Indonesia the black band runs between the rhinophores and bifur-
cates and reconnects with the primary black band that encircles most of the notum. In the Indonesian
specimens an additional black band connects the primary oval of black anterior to the branchial
plume. All four specimens have unipinnate branchial leaves, a single row of tubercles around the
branchial sheath and a pair of tubercles around the rhinophore sheath. One of the Papuan specimens
has one rhinophoral tubercle that is the same color as the rhinophore while the other tubercles is
opaque white. In the remaining three specimens, both rhinophoral tubercles are opaque white. It is ev-
ident that all four specimens discussed here, likely represent a single species, A. williamsi.

Despite some variability in the external anatomy of both A. albatrossae and A. williamsi, there are
consistent differences that clearly separate them. In A. albatrossae, there is always a T- shaped mark in
front and between the rhinophores and then bifurcates to form a black oval that terminates at the ante-
rior end of the branchial sheath, while in A. wil/iamsi there is a continuous black oval band that extends
from the anterior end of the notum and encircles the posterior end of the branchial sheath. Yel-
low-orange pigment is present in all specimens of A. al/batrossae, while in A. williamsi, none of this
pigment is present. In A. albatrossae, there is a double row of tubercles around the branchial sheath
while in A. williamsi there is only a single row. In A. albatrossae, there are four tubercles around the
rhinophoral sheath while in A. williamsi there are only two tubercles. Aldisa albatrossae consistently
has more tubercles (4-5 rows outside of the oval band and 4—5 additional rows inside the band) than
does A. williamsi (2-3 rows outside of the oval band and 2-3 additional rows inside the band). The gill

ELWOOD, VALDES, AND GOSLINER: ALDISA 179

FiGuRE 5. Aldisa williamsi, sp. nov. (CASIZ 109791). A. Reproductive system, scale bar = 0.5 mm; B. Penial hooks, scale
bar = 5 pm; C. Ventral view of the anterior border of the foot, scale bar = 1 mm. Abbreviations: am, ampulla; be, bursa
copulatrix; dd, deferent duct; fg, female gland; pr, prostate; sr, seminal receptacle; v, vagina.

180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 14

branches of A. albatrossae appear to be more numerous (6-8) than those of A. williamsi (6) and more
highly pinnate.

The radular teeth of both species are similar in that there is a pair of hamate teeth in addition to the
elongate pectinate teeth. In A. albatrossae there are 20—24 denticles along the pectinate teeth while in
A. williamsi there appear to be fewer (15-20).

A few differences are seen between the reproductive systems of the two species. In one specimen
of A. williamsi the seminal receptacle is about a quarter the size of the bursa copulatrix, while the semi-
nal receptacle in one specimen of A. albatrossae almost equals the size of its bursa copulatrix. The
uterine tube of A. williamsi connects to the female gland at a point much closer to the genital atrium
than does the uterine tube of A. al/batrossae. These reproductive and radular differences should be re-
confirmed when additional material of A. williamsi becomes available.

There are consistent differences in the external morphology of these two species that warrant
their description as distinct taxa. The fact that these differences are consistent within overlapping geo-
graphical ranges in Indonesia is supportive of the fact that they do not simply represent geographical
variants of a single species.

Discodoris liturata Bergh, 1905, is another dorid with a similar color pattern. It has a gray-green
body with opaque white pustules and black markings. The radular morphology of this species has typ-
ically hamate teeth and lacks any of the pectinate teeth of A/disa (Rudman 1998c). Both D. liturata
and A. williamsi bear a strong resemblance to another dorid, Phyllidiella pustulosa (Cuvier, 1804),
and are probably mimics of this species (Rudman 1998b; 2000). In P. pustulosa a dorsal gill and a
radula are absent. Phyllidiella pustulosa and D. liturata also differ from A. williamsi and A.
albatrossae in having black rather than off-white to pale brown rhinophores.

Two undescribed species, referred to as Doris? sp. 2 (Rudman 1998a) and Chromodoris sp.
(Debelius 1996:213, bottom photo) are similar in their appearance to this group of Phyllidia mimics.
Doris? sp. 2, from Thailand, is quite similarly colored to 4. albatrossae, but it has consistent differ-
ences in its color pattern. It has black rather than off white to pale brown rhinophores and has yellow
pigment on the tubercles rather than between them. It also has a complete oval of black pigment anteri-
orly rather than a T-shaped mark. Also the black oval terminates posterior to the branchial sheath
rather than anteriorly. Chromodoris sp. of Debelius is blue in color with irregular black markings and
yellow pustules on the notum that are similar in appearance to Phyllidia varicosa Lamarck, 1801. We
were unable to obtain specimens to examine radular or reproductive systems characters. Therefore it
is not possible, at present, to state anything further about their systematic placement.

ACKNOWLEDGMENTS

The material studied has been collected by Gary Williams and Robert Bolland. Jim Black kindly
provided a photo of Aldisa albotrossae from Indonesia. This paper was supported by the National Sci-
ence Foundation through the PEET Grant DEB - 9978155 (Phylogenetic systematics of dorid nudi-
branchs) and the REU Grant DBI - 9820251 (Summer Systematics Institute).

LITERATURE CITED

BERTSCH, H. AND S. JOHNSON. 1982. Three new species of dorid nudibranchs from the Hawaiian Islands. The
Veliger 24:208-218.

DEBELIUS, H. 1996. Nudibranchs and sea snails, Indo-Pacific field guide. IKAN-Unterwasserarchiv, Frankfurt,
321 pp.

MASuDA, H. 1999. Guide Book to Marine Life. Tokai University Press, Tokyo, 404 pp.

MILLEN, S. V. AND T. M. GOSLINER. 1985. Four new species of dorid nudibranchs belonging to the genus Aldisa,
with a revision of the genus. Zoological Journal of the Linnean Society 84:195—233.

Ono, A. 1999. Opisthobranchs of Kerama Islands. TBS-Britannica, Tokyo, 110 pp.

ELWOOD, VALDES, AND GOSLINER: ALDISA 181

ORTEA, J. AND M. BALLESTEROS. 1988. Descripcion de una espectacular especie del género Aldisa Bergh, 1878.
Bollettino Malacologico 24:155—160.
RUDMAN, W. B. 1998a. Doris? sp. 2 available via http://www.seaslugforum.net/dorisp2.htm.
. 1998b. Mimicry—Phyllidiella, flatworms, Chromodoris available via http://www.seaslugforum.
net/mimicry.htm.
1998c. Discodoris? liturata Bergh, 1905 available via http://www.seaslugforum.net/disclitu.htm
2000. Doris? sp. 10 available via http://www.seaslugforum.net/dorisp10.htm.

© CALIFORNIA ACADEMY OF SCIENCES, 2000
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San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
ious Library |
Volume 52, No. 15, pp. 183-193, 7 figs., 1 table. 1 August 21, 2001

| SEP 05 200t |

The Identity of Doris (s.1.) Species MacFarland, 1966
(Mollusca, Nudibranchia, Discodorididae): A Persistent Mystery
from California Solved

by
David W. Behrens and Angel Valdés!

Department of Invertebrate Zoology and Geology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

The Californian dorid nudibranch species, previously referred to by several authors as Do-
ris (s.l.) species, based upon its original reference in MacFarland (1966), is a synonym of
Diaulula sandiegensis (Cooper, 1863). The single specimen examined by MacFarland, and
other animals collected from southern California, matching the external coloration of Do-
ris (s.L.) species, have been examined and their internal anatomy is identical to that of
Diaulula sandiegensis. Some variation has been observed in the shape of the outermost
radular teeth of this species. The obscure and poorly described species Doris odonoghuei
Steinberg, 1963 (= Doris echinata O’ Donoghue, 1922) is probably a synonym of Diaulula
sandiegensis as well.

The monographic work “Studies of the Opisthobranchiate Mollusks of the Pacific Coast of North
America” by F. M. MacFarland was published posthumously in 1966. This work is composed of a se-
ries of unpublished notes that MacFarland had been preparing at the time of his death. One of several
of the undescribed species included was referred to as Doris (s./.) species. MacFarland frequently
used Latin abbreviations in his notes and manuscripts, and in this instance “s.|.” referred to the Latin,
sensu lato, in the broader sense. MacFarland (1966) did not provide a specific name for this animal,
which has similar external morphology and coloration to Diaulula sandiegensis (Cooper, 1863). He
examined only one specimen of Doris (s./.) species (Fig. 1), collected from Arch Rock Pool, Newport
Bay, California, but he never studied it anatomically. The name Doris (s./.) has been carried in the lit-
erature in numerous publications (Sphon and Lance 1968; Behrens 1980; McDonald and Nybakken
1981; McDonald 1983), and it is normally used for dorid nudibranchs similar to Diaulula
sandiegensis but having pale dorsal spots. Other authors referred to this animal as Doris sp. (McDon-
ald and Nybakken 1981; McDonald 1983) or Diaulula sp. | (Behrens 1991, 1992). All these refer-
ences are based on MacFarland’s descriptions and newly collected specimens as well. However, the
question of whether this animal constitutes a different species from Diaulula sandiegensis remains
unresolved.

Prior to the publication of MacFarland’s (1966) memoir, a species with similar external charac-
teristics was described as Doris echinata by O’ Donoghue (1922). Later, Iredale and O’ Donoghue
(1923) reassigned this species to Doridigitata d’Orbigny, 1839 and changed the name (without expla-
nation) to Doridigitata maculata. Steinberg (1963) noted that both O’ Donoghue’s names were preoc-
cupied by Doris echinata Lovén 1846 and Doris maculata Garstang 1896, respectively, and proposed
a new name, Doris odonoghuei for this species. Additionally, Steinberg (1963) questioned whether

|Current address: Department of Malacology, Museum of Natural History of Los Angeles County, 900 Exposition Boule-
vard, Los Angeles, California 90007.

183

184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 15

the placement of this species in the genus Doris is accurate and suggested that further study was neces-
sary.
In this paper we attempt to determine the identity of Doris (s./.) species based on the study of
MacFarland’s original material and additional specimens deposited at the Department of Invertebrate
Zoology and Geology of the California Academy of Sciences (CASIZ). In addition, the status of Doris
odonoghuei is discussed.

MATERIAL AND METHODS

For this paper several species matching the external coloration described for Diaulula
sandiegensis and Doris (s.l.) species were examined. Table | summarizes the material sources and
collection localities. Specimens were dissected by dorsal incision. Their internal features were exam-
ined and drawn under a dissecting microscope using a camera lucida. Parts of the dorsum have been
critical point dried for scanning electron microscopy (SEM) of the caryophyllidia. Special attention
was paid to the morphology of the reproductive system and digestive system, including the radulae,
which have been prepared for examination on SEM. Features of living animals were recorded from
photographs or notes of collectors.

DESCRIPTIONS

EXTERNAL MORPHOLOGY.—The living animals measured up to 53 mm in length. The back-
ground color varies from translucent white to tan (Figs. 1; 2A, B). The notal surface is covered with
brown specks and bears numerous irregular oval brown spots. In most specimens the center of the
brown spots is lighter in color, surrounded by a dark ring. A white band, composed of minute white
specks occurs along the notal margin. The body is oval, highest along its midline, sloping gradually to
the margins. The notal surface is densely covered with various sized caryophyllidia (Fig. 3D). The gill
is completely retractile into a branchial pit. The six tripinnate branchial leaves are upstanding and do
not spread to the edges of the notum. The branchial leaves are white to cream in color and are densely
sprinkled with brown specks. The anal papilla is located at the center of the branchial plume. The
rhinophores are perfoliate with 12—18 lamellae and are retractile into short upright sheaths. The color
of the rhinophores is similar to that of the branchial plume. Ventrally, the foot is grooved and notched,
wide, tapering posteriorly into a round end. The posterior end of the foot extends only slightly beyond
the posterior margin of the notum. The oral tentacles are slender and pointed distally (Fig. SC).

ANATOMY.—The labial cuticle is smooth. The radular formula is 14 x 16.0.16 in a 10-mm-long
specimen (CASIZ 060977), 15 x 23.0.23 in an 18-mm-long specimen (CASIZ 025880) and
22 x 27.0.27 in a 46-mm-long specimen (CASIZ 068277). There is no trace of rachidian teeth. The
lateral teeth (Figs. 3A, B; 4A, B) are simple hamate increasing in size from the center of the radular
ribbon to the ninth and tenth tooth, then decreasing to the margin. The outermost two lateral teeth are
very elongate and are smooth (Fig. 3C), or have one to three small denticles (Fig. 4C, D), depending
on the specimen.

The reproductive system is triaulic (Fig. SA, B). The ampulla is tubular and convoluted. It nar-
rows into a short thin tube and connects to the oviduct and prostatic portion of the vas deferens. Imme-
diately after branching, the oviduct enters the massive female gland mass. The vas deferens is long
and slightly thinner than the ampulla, until it expands into two wide and large, contiguous prostatic
portions. A long, thin duct emerges from the prostatic portion and becomes highly convoluted in the
ejaculatory segment, prior to entering a common genital atrium with the vagina. The vaginal duct is
thick, normally straight and connects to the large, round bursa copulatrix. A separate duct from the
bursa copulatrix connects to the smaller, spherical seminal receptacle. A short, thin uterine tube

BEHRENS AND VALDES: IDENTITY OF DORIS (SL.) 185

FiGurE 1. Specimen identified by MacFarland (1966) as Doris (s./.) species (CASIZ 025880). A. Photograph of the living
animal taken by G. E. MacGinitie. B, C. Drawings published by MacFarland (1966, pl. 25, figs. 1, 2).

emerges near the connection of this duct and then connects the seminal receptacle to the female gland
near the genital atrium.

DISCUSSION

After the anatomical study of the material of Doris (s./.) species examined by MacFarland
(CASIZ 025880), of additional specimens with a similar external coloration from southern California
(CASIZ 060976), and of specimens matching the original description of Diaulula sandiegensis
(CASIZ 068277; CASIZ 071641), we were unable to find any consistent differences. It is clear that
Doris (s.1.) species constitutes a color variation of Diaulula sandiegensis. The external coloration of
this species is extremely variable. It ranges from white or cream to yellow, with brown rings or solid
spots, sometimes surrounded by an opaque white ring (Fig. 2). Specimens from Canada, Alaska, and
the Russian far east generally have the dorsum covered with numerous very dark spots (Fig. 2E, F),
whereas in southern California and Mexico the spots are lighter and less common. Specimens from
central and northern California, Oregon, and Washington match the original description by Cooper
(1863).

The reproductive system of all the specimens examined has two large and distinct prostatic re-
gions in the vas deferens. There is a long, thick, straight vaginal duct. The oviduct, vas deferens and
uterine duct all enter the female gland mass in the same proximity, near the genital atrium (Figs. 5A,
B; 6A, B).

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

186

Volume 52, No. 15

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BEHRENS AND VALDES: IDENTITY OF DORIS (S.L.) 187

FIGURE 2. Living animals of Diaulula sandiegensis (Cooper, 1863). A. Specimen from San Diego, California, originally
identified as Doris (s./.) species; B. Specimen from Orange County, California, originally identified as Doris (s./.) species; C.
Specimen from San Luis Obispo, California; D. Specimen from the Channel Islands, California; E. Specimen from Vancouver
Island, British Columbia, Canada; F. Specimen from Vancouver Island, British Columbia, Canada. Photograph 1A by J.
Hamann, others by D. Behrens.

Camera lucida drawings based on light microscopy of the radula of Doris (s./.) species
(MacFarland 1966; McDonald 1983, 1997; Behrens 1992) suggests that the species has smooth,
hamate, outer lateral teeth. Scanning electron microscopy of the specimen seen by MacFarland (Fig.
4C, D) shows the presence of denticles on the outer two lateral teeth. This character is not present in
other examined specimens with the same color pattern collected from southern California (Fig.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 15

FIGURE 3. Diaulula sandiegensis (Cooper, 1863), scanning electron micrographs of a specimen originally identified as Doris
(s./.) species (CASIZ 060977). A. Inner lateral teeth; B. Lateral teeth from central portion of half-row; C. Outer lateral teeth;
D. Caryophyllidia.

BEHRENS AND VALDES: IDENTITY OF DORIS (SL.) 189

FiGurE 4. Diaulula sandiegensis (Cooper, 1863), scanning electron micrographs of a specimen identified by MacFarland
(1966) as Doris (s.1.) species (CASIZ 025880). A. Inner lateral teeth; B. Lateral teeth from central portion of half-row; C. Outer
lateral teeth; D. Detail of the denticles on the outer lateral teeth.

190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 15

B

am

bc

FiGurE 5. Diaulula sandiegensis (Cooper, 1863), anatomy of a specimen originally identified as Doris (s.1.) species (CASIZ
060977). A. Reproductive system, scale bar = 1 mm; B. Detail of several organs, scale bar = 1 mm; C. Ventral view of the
mouth area, scale bar = 1 mm. Abbreviations: am, ampulla; bc, bursa copulatrix; dd, deferent duct; fg, female gland mass; ot,
oral tentacle; pr1, proximal prostatic region; pr2, distal prostatic region; sr, seminal receptacle; v, vagina.

3A-C), so it is clearly due to intraspecific variation. Other specimens of Diaulula sandiegensis have
smooth, sharply-pointed, hamate-shaped teeth across the entire row (Fig. 7A—C), identical to those of
Doris (s.1.) species.

Scanning electron microscopy of Diaulula sandiegensis clearly indicates the presence of
caryophyllidia (Fig. 7D), which are identical in size and density to those present in Doris (s./.) species
(Fig. 3D).

O’Donoghue’s (1922) description of Doris echinata was brief, stating simply that the dorsum is
covered with spiculate papillae and the color is opaque white with from a dozen to forty small brown
spots scattered irregularly over the surface. The radula was described as simply hamate, 16—18 rows
of 13-15 lateral teeth per half-row. A description of the reproductive system is lacking, except for
mention that the penis is unarmed. O’ Donoghue (1922) stated that though he felt that the classification
of the family was unsatisfactory, Doris echinata falls within its definition.

In proposing the name Doris odonoghuei to rectify the preoccupancy issue discussed earlier,
Steinberg (1963) examined two specimens from the collection of the Friday Harbor Marine Labora-
tories. Questioning the assignment of the species to Doris, she dissected the smaller of the two, but
came to no satisfactory conclusion. Recent review of her personal notes (J. Steinberg, pers. commun.,
Jan. 2001) revealed no further evidence to assist in its placement.

BEHRENS AND VALDES: IDENTITY OF DORIS (S.L.) 19]

|

FiGurE 6. Diaulula sandiegensis (Cooper, 1863), anatomy (CASIZ 068277). A. Reproductive system, scale bar = 1 mm; B.
Detail of several organs, scale bar = 1 mm; C. Ventral view of the mouth area, scale bar = 1 mm. Abbreviations: am, ampulla;
bc, bursa copulatrix; dd, deferent duct; fg, female gland mass; ot, oral tentacle; pr1, proximal prostatic region; pr2, disstal pros-
tatic region; sr, seminal receptacle; v, vagina.

Since that time no published accounts or casual observations have been made of this species. San-
dra Millen (pers. commun., April 1982) indicated that she had never collected specimens in the Van-
couver area, British Columbia, that she could clearly identify as Doris odonoghuei. According to
Millen it is impossible to distinguish Doris odonoghuei from small Diaulula sandiegensis.

Whereas this species has not been definitely confirmed since O’ Donoghue’s (1922) original de-
scription and no type material is available for examination, and whereas this description cannot be dif-
ferentiated from Diaulula sandiegensis, we propose that this species be regarded as a synonym of
Diaulula sandiegensis.

ACKNOWLEDGMENTS

We thank Jeff Goddard for collection of material, Jeff Hamann for use of his photograph and Joan
Steinberg and Sandra Millen for sharing their field and laboratory notes. Joan Steinberg made con-
structive comments on the manuscript. This paper has been supported by the National Science Foun-
dation, through the PEET grant (DEB-9978 155, “Phylogenetic systematics of dorid nudibranchs”) to
Terrence M. Gosliner.

192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 7. Diaulula sandiegensis (Cooper, 1863), scanning electron micrographs (CASIZ 068277). A. Inner lateral teeth; B.
Lateral teeth from central portion of half-row; C. Outer lateral teeth; D. Caryophyllidia.

BEHRENS AND VALDES: IDENTITY OF DORIS (SL.) 193

LITERATURE CITED

BEHRENS, D. W. 1980. Pacific coast nudibranchs: A guide to the opisthobranchs of the northeastern Pacific. Sea
Challengers, Los Osos, California. 112 pp.

. 1991. Pacific coast nudibranchs: A guide to the opisthobranchs of the northeastern Pacific, 2nd ed. Sea

Challengers, Monterey, California. 107 pp.

. 1992. Pacific coast nudibranchs. Supplement I — Radula. Sea Challengers, Monterey, California. 11 pp.

BERGH, R. 1880. On the nudibranchiate gastropod Mollusca of the North Pacific Ocean, with special reference to
those of Alaska. Scientific Results of the Exploration of Alaska 1(Art. 6):189-276, pls. 9-16.

CoopPeER, J. G. 1863. Some genera and species of California Mollusca. Proceedings of the California Academy of
Natural Sciences 2:202—207.

IREDALE, T. AND C. H. O'DONOGHUE. 1923. List of British nudibranchiate Mollusca. Proceedings of the
Malacological Society of London 15:195—233.

MACFARLAND, F. M. 1966. Studies of the Opisthobranchiate Mollusks of the Pacific coast of North America.
Memoirs of the California Academy of Sciences, No. 6. 546 pp., 71 pls.

Marcus, ER. 1961. Opisthobranch mollusks from California. Veliger 3(Supp):1—85.

MCDONALD, G. R. 1983. A review of the nudibranchs of the California coast. Malacologia 24:1 14-276.

. 1997. A review of the nudibranchs of the California coast. Master’s thesis, California State University,
Hayward. 337 pp.

MCDONALD, G. R. AND J. W. NYBAKKEN. 1981. Guide to the nudibranchs of California, 2nd ed. American
Malacologists Inc., Melbourne, Florida. 72 pp.

O’ DONOGHUE, C. H. 1922. Notes on the Nudibranchiate Mollusca from the Vancouver island region. III. Re-
cords of species and distribution. Transactions of the Royal Canadian Institute 14:145—167.

SPHON, JR.,G. G. AND J. R. LANCE. 1968. An annotated list of the nudibranchs and their allies from Santa Barbara
County, California. Proceedings of the California Academy of Sciences 36:73—84.

STEINBERG, J. E. 1963. Notes on the opisthobranchs of the west coast of North America — III. Further nomencla-
tural changes in the order Nudibranchia. Veliger 6:63-67.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

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Volume 52, No. 16, pp. 195-208, 10 figs. August 21, 2001

| |
A New Species of the Soft Coral Genus Eleutherobi
(Octocorallia: Alcyoniidae) from South?

by

Gary C. Williams and Susan A. Little
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Goiden Gate Park, San Francisco, California 94118

Putter, 1900,
rica

The alcyoniid soft coral genus Eleutherobia was previously known to contain sixteen valid
species, distributed from southern Africa to the south central Pacific Ocean. A new species
is here described from the KwaZulu-Natal coast of South Africa, thereby making a total of
seventeen species, with four of these restricted to southern Africa. The new taxon is superfi-
cially similar to Eleutherobia rubra (Brundin, 1896), originally described from Japan, but
differs from it and all other species of the genus by a unique complement of sclerite types.

Williams (2000b:159) summarized recent discoveries concerning natural products biochemistry
and the soft coral genus E/eutherobia. In the same paper, Eleutherobia zanahoria was described from
the Tonga Islands, thus extending the known geographic range of the genus approximately 3000 km
to the southeast into the central South Pacific. Four species of the genus are known from southern Af-
rica, including the new species described below. The other taxa are Eleutherobia studeri (J. S.
Thomson, 1910), E. rotifera (J. S. Thomson, 1910), and E. aurea Benayahu and Schleyer, 1995 (Wil-
liams 1992a, 2000a, 2000b). Together with thirteen species from the Indo- West Pacific, a total of sev-
enteen species of the genus Eleutherobia are presently considered valid. The geographic range of the
genus extends from the southeastern fringe of the Atlantic Ocean (Cape Peninsula), around South Af-
rica, through the Indian Ocean, and as far as Japan, Palau, and Tonga in the Pacific Ocean (Fig. 1). A
revision, which will add other taxa for southern Africa, is currently in progress by the authors.

METHODS

Material was collected by SCUBA or dredge and preserved in 70% ethanol. Sclerites were iso-
lated using sodium hypochlorite (household bleach). Micrographs and photographs for Figures 2 and
3 were taken using a Nikon Coolpix 990 digital camera, a Nikon SMZ-10 dissecting microscope, and
an Olympus CH-2 compound microscope. Scanning electron micrographs were taken using a Leo
1400 Series scanning electron microscope. Sclerites were examined and drawn using an Olympus
CH-2 compound microscope with an attached drawing tube. Digital images and plates of photo-
graphs, micrographs, and scanning electron micrographs were made using Adobe Photoshop soft-
ware. The abbreviation used for the South African Museum, Cape Town, is SAM.

195

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|_|
Durban. re

East London,

Port Elizabeth, Algoa Bay
Cape St. Francis

sCape Town

Cape

Peninsula C@Pe Aguthas

FiGURE |. A. Map of southern Africa showing collecting stations for Eleutherobia vinadigitaria sp. nov. B. Map of the
Indo-West Pacific showing geographic distribution of the genus Eleutherobia. Arrows show type locality of Eleutherobia
vinadigitaria sp. nov. Numbers along axes represent degrees of longitude and latitude.

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA 197

FIGURE 2. Eleutherobia vinadigitaria sp. nov. A. Holotype, total length 97 mm. B. Paratype exterior, total length 88 mm.
C. Paratype interior, longitudinal section; total length 88 mm. D. Holotype, detail of surface of polyparium showing re-
tracted polyps; scale bar = 0.50 mm. E. Holotype, sclerites from the surface of the polyparium; scale bar = 0.07 mm.

198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SYSTEMATIC ACCOUNT

Family Alcyoniidae Lamouroux, 1812
Eleutherobia Pitter, 1900

Eleutherobia Piitter, 1900:449. Verseveldt and Bayer, 1988:27. Williams, 1992a:306. 2000b:160.
Nidalia (in part): non Gray, 1835.

Bellonella (in part): non Gray, 1862.

Metalcyonium (in part): non Pfeffer, 1889.

DIAGNOSIS.—A Icyoniid soft corals with colonies digitiform, finger-shaped, often conical and ta-
pering, rarely multilobate. Polyps monomorphic. Calyces absent, although retracted polyps often
form low rounded to conspicuous protuberances. Sclerites mostly derived from radiates although
spindles, crosses, barrels, or tuberculate spheroids may also be present. Polyp sclerites often present
as eight points, or crown and points, or totally absent. Zooxanthellae absent.

TYPE SPECIES.—Eleutherobia japonica Piitter, 1900, by monotypy.

DIVERSITY AND DISTRIBUTION.—Seventeen species of the Indo-West Pacific (southern and east-
ern Africa to Japan and Tonga).

Eleutherobia vinadigitaria sp. nov.
Figs. 1-10

MATERIAL EXAMINED.—HOLOTYPE: SAM-H4877, station number DEEP No. 1, South Africa,
KwaZulu-Natal, outer anchorage off Durban Bluff, 52 m depth, 13—14 December 1984, collected by
W. R. Liltved with aid of SCUBA, one whole specimen (97 mm in length). PARATYPE: SAM-H4878,
same data as holotype, one specimen (cut longitudinally into two halves; each half 88 mm in length).
OTHER MATERIAL: SAM-H792, station number P. F. 11538, South Africa, KwaZulu-Natal, Thukela
(Tugela) River Mouth, NW by N 22 1/2 miles, 86 m depth, 29 January 1901, collected by S.S. Pieter
Faure survey with aid of dredge, one whole specimen (20 mm in length). SAM H-4835, same data as
holotype, three whole specimens (18 mm, 72 mm, and 100 mm in length); the latter specimen partly
cut transversely, thus partially separating the polypary from the stalk (Fig. 4B).

DIAGNOSIS.—A|cyoniid soft corals with digitiform colony shape, tapering distally to a rounded
apex. Stalk conspicuous, comprising 20 to 50% of total colony length. Retracted polyps form
moundlike protuberances. Sclerites of the eight polyp wall ridges with longitudinal rows of mostly
sparsely-set, elongated, needle-like spindles (up to 0.30 mm long), sometimes arranged more or less
in a chevroned fashion. Coenenchymal sclerites restricted to a thin surface layer of the polypary and
stalk, primarily radiates and spindles, with a few clubs also present (0.03—0.20 mm in length). Interior
sclerites absent. Color of stalk pink; polypary pink, red, or deep wine red. Sclerite color primarily red-
dish, rarely colorless.

DESCRIPTION OF THE HOLOTYPE.—Growth form and size. The wet-preserved holotype is
97 mm in length, and varies from 10 mm in width near the distal tip to 25 mm in width at the base of
the stalk. It is finger-shaped, not branched or lobate. The length of the polypary is 77 mm, and com-
prises 79% of the total colony length, while the stalk is approximately 20 mm long and represents
about 21% of the total colony length. The specimen is digitiform and tapers markedly from the
holdfast to the distal terminus, which is conspicuously rounded (Fig. 2A).

Polyps. The polyps are tightly retracted in the available specimens, including the holotype. A
small portion of the surface coenenchyme of the holotype was cut away to expose the retracted polyps.
Many of the exposed polyps contain several mature gonads in their gastric cavities. These are pale yel-
low in color and mostly vary from 0.2 — 0.4 mm in diameter. The size of retracted polyps on the sur-

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA 199

A

FiGure 3. Eleutherobia vinadigitaria sp. nov. Living soft corals. A. Several colonies with polyps retracted (Photograph by
Michael Schleyer). B. A single colony with polyps expanded (Photograph by Michael Schleyer).

200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Ficure 4. Eleutherobia vinadigitaria sp. nov. Micrographs of surface region of paratype; longitudinal section of polypary.
A. A single retracted polyp showing chevroned arrangement of needle-like sclerites in polyp wall, and thin layer of sclerites of
mostly radiates in the surface coenenchyme. B. Three adjacent polyps. Scale bars each represent 1.0 mm. Abbreviations: cs —
coenenchymal sclerites, gc — gastric cavity, ic — interior coenenchyme, p — points of enchevroned needle-like sclerites, rp — re-
tracted polyp, sc — surface coenenchyme.

face of the polypary varies from 0.3 to 0.8 mm in diameter (Fig. 2D). Observation of living non-type
material at the type locality shows that the polyps are generally retracted in bright daylight (Fig. 3A)
and are expanded in early hours of the morning (Fig. 3B) (M. H. Schleyer, pers. comm.). Polyp scler-
ites are arranged in eight longitudinal rows along the ridges of the polyp walls, relatively sparsely
placed. In some polyps, the sclerites may be disposed in a chevroned fashion, forming eight points.

Sclerites. Sclerites from the polyp body walls are needle-like spindles, mostly with tuberculation
restricted to the opposite margins, as the flattened faces of the sclerites are for the most part smooth
and devoid of ornamentation (Figs. 6A, 7A-B, 8A). They vary in length from 0.17 to 0.30 mm. Scler-
ites are lacking in the tentacles and pinnules. Coenenchymal sclerites are restricted to a very thin sur-
face layer of the polypary and stalk (Fig. 4B). Sclerites of the surface of the polypary, as well as the
bases of the polyps, are radiates and spindles (0.03—0.20 mm in length). Some of the spindles may be
somewhat club-shaped (Fig. 6B, 10B). Sclerites from the surface of the stalk and holdfast are radiates
(0.05—0.11 mm in length). Sclerites are altogether absent from the interior of the colonies.

Color. The interior of the wet-preserved holotype is cream colored to pale yellow (Fig. 2C), but
crisp white in life (M. H. Schleyer, pers. commun.). The polyps are white to cream white (Fig. 3B).
The color of the coenenchyma! sclerites varies from deep red (Fig. 2E) to pale red, while the polyp
sclerites vary from pale red to colorless.

INTERNAL ANATOMY OF THE PARATYPE.—The paratype specimen was cut longitudinally to re-
veal aspects of internal anatomy (Figs. 2C; 4A, B). The mature polyps have gastric cavities that extend
throughout the entire length of the colony (Fig. 2C). The surface coenenchyme containing sclerites is

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA 201

Ficure 5. Eleutherobia vinadigitaria sp. nov. Variation in colony shape and size (non type material, wet preserved).
A. SAM-H4835, 72 mm length. B. SAM-H4835, 100 mm. C. SAM-H4835, 18mm. D. SAM-H792, 20 mm. Scale
bar = 20 mm.

very thin, mostly < 0.2 mm in thickness (Fig. 4). The interior coenenchyme is firm but lacks sclerites
(Fig. 4B). Conspicuous points formed by enchevroned sclerites from the polyp walls of the
anthocodial neck zones can clearly be observed. However, transversely disposed sclerites forming a
distinctive crown are lacking or at most are very sparsely distributed below the points (Fig. 4A).

ETYMOLOGY.—The specific epithet of the new species is derived from the Latin, vinum (wine),
digitus (a finger), and the suffix, -aria (like); in reference to the wine red, finger-shaped colonies of
this species of soft coral.

DISTRIBUTION.—The new species is known only from two localities in KwaZulu-Natal, South
Africa (Fig. 1).

VARIATION.—The six available specimens range in size from 18-100 mm in length. The
polypary comprises approximately 50-80% of the total colony length (Figs. 2, 5). The stalk and the
polypary are approximately equal in length in the smallest specimens (Fig. 5C, D).

DISCUSSION

The rachises of the larger specimens of Eleutherobia vinadigitaria sp. nov. have retracted polyps
exhibiting a variety of diameters (Fig. 2A, B, D). These all appear to be autozooids in various interme-
diate states of growth—each with eight equal-sized lappets closing over the opening of the retracted
polyps, anthocodiae, and well-developed gastric cavities (Figs. 2C, 4B). In dimorphic taxa such as the

202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 6. Eleutherobia vinadigitaria sp. nov., holotype. Variation in sclerite shape and size. A. Sclerites from the polyp
walls. B. Coenenchymal sclerites from the surface of the polypary. C. Coenenchymal sclerites from the surface of the holdfast
region of the stalk. Scale bar = 0.10 mm.

various species of the superficially similar soft coral genus Paraminabea Williams and Alderslade,
1999, the siphonozooids and autozooids display two distinct sizes without intermediates (Williams
1992b; Williams and Alderslade 1999). An additional aspect of distinction between the two genera is
as follows: Eleutherobia vinadigitaria sp. nov., as in most other species of the genus, has distinctive
longitudinal rows of needle-like sclerites in the neck zone of the anthocodiae (often enchevroned),
while all species of Paraminabea apparently lack polyp sclerites (Fabricius and Alderslade, in press;
Williams 1992b; Williams and Alderslade 1999).

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA

FIGURE 7. Eleutherobia vinadigitaria sp. nov. Scanning electron micrographs of sclerites from the holotype. A-B. Sclerites
from the polyp wails, each 0.27 mm. C-K. Coenenchymal sclerites from the surface of the polypary and polyp bases. C.
0.09 mm. D. 0.07 mm. E. 0.05 mm. F. 0.06 mm. G. 0.05 mm. H. 0.08 mm. I. 0.09 mm. J. 0.06 mm. K. 0.08 mm.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 16

FIGURE 8. Eleutherobia vinadigitaria sp. nov. Scanning electron micrographs of sclerites from the holotype. A. A polyp wall
sclerite, 0.20 mm. B—G. Coenenchymal sclerites from the surface of the polypary and polyp bases. B. 0.04 mm. C. 0.04 mm.
D. 0.06 mm. E. 0.05 mm. F. 0.05 mm. G. 0.07 mm.

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA

FiGuRE 9. Eleutherobia vinadigitaria sp. nov. Scanning electron micrographs of coenenchymal sclerites from the surface of
the stalk of the holotype. A. 0.05 mm. B. 0.01 mm. C. 0.10 mm. D. 0.03 mm. E. 0.09 mm. F. 0.05 mm.

206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 10. Eleutherobia vinadigitaria sp. nov. Scanning electron micrographs of coenenchymal sclerites from the surface of
the stalk of the holotype. A. 0.07 mm. B. 0.06 mm. C. 0.06 mm. D. 0.09 mm.

WILLIAMS: ELEUTHEROBIA FROM SOUTH AFRICA 207

Eleutherobia vinadigitaria sp. nov. superficially resembles Eleutherobia rubra (Brundin, 1896)
from Japan and northwestern Australia (compare Figs. | and 3 with Verseveldt and Bayer 1988, fig.
32e-h). However, Eleutherobia rubra can be differentiated from the new species by the following
characters: well-developed crown and points with densely set sclerites in the anthocodiae, sclerites in
the tentacles, and highly ornamented polyp wall spindles with elongated tubercles (Verseveldt and
Bayer 1988, figs. 30-31). Eleutherobia vinadigitaria sp. nov., on the other hand, has weakly devel-
oped points (without the development of a crown), lacks tentacle sclerites, and has polyp wall sclerites
that are sparsely ornamented spindles with low tubercles.

Eleutherobia aurea Benayahu and Schleyer (1995), also from Natal, South Africa, is similar in
colony shape to Eleutherobia vinadigitaria sp. nov., but differs by having bright yellow coloration, a
lack of polyp sclerites, and coenenchymal sclerites that are compact radiates and spheroids.

ACKNOWLEDGMENTS

We express our gratitude to Yehuda Benayahu (Tel Aviv University, Tel Aviv), Michael
Schleyer (Oceanographic Research Institute, Durban), Leen van Ofwegen (National Museum of Nat-
ural History, Leiden), Phil Alderslade (Museum and Art Gallery of the Northern Territory, Darwin),
and Alan Leviton, Katie Martin, and Marilyn Eversole (California Academy of Sciences, San Fran-
cisco) for their support, suggestions, and comments.

This paper is a result of a collaborative effort during the 1997 (June through August) Summer
Systematics Institute, California Academy of Sciences, San Francisco.

LITERATURE CITED

BENAYAHU, Y. AND M. H. SCHLEYER. 1995. Corals of the south-west Indian Ocean II. Eleutherobia aurea spec.
nov. (Cnidaria, Alcyonaria) from deep reefs on the KwaZulu-Natal Coast, South Africa. South African As-
sociation for Marine Biological Research, Oceanographic Research Institute, Investigational Report
68:1—12.

FABRICIUS, K. AND P. ALDERSLADE. In press. Soft corals and sea fans—a comprehensive guide to the tropical
shallow water genera of the central-west Pacific, the Indian Ocean and the Red Sea. Australian Institute of
Marine Science, Townsville.

GRAY, J. E. 1835. Characters of a new genus of corals (Nidalia). Proceedings of the Zoological Society of Lon-
don 3:59-60.

. 1862. Description of two new genera of zoophytes (Solenocaulon and Bellonella) discovered on the
north coast of Australia by Mr. Rayner. Proceedings of the Zoological Society of London 1862:34—37.

PFEFFER, G. 1889. Zur Fauna von Sud-Georgien. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten
6(2):49-55.

PUTTER, A. 1900. Alcyonaceen des Breslauer Museum. Zoologische Jahrbiicher (Systematik) | 3(5):443—462.

THOMSON, J. A. AND L. M. I. DEAN. 1931. The Alcyonacea of the Siboga Expedition with an addendum to the
Gorgonacea. Siboga Expedition Monographs 13d:1—227.

THOMSON, J. S. 1910. The Alcyonaria of the Cape of Good Hope and Natal. Alcyonacea. Transactions of the
Royal Society of Edinburgh 47(3):549-589.

VERSEVELDT, J. AND F. M. BAYER. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and
Nidaliopsis (Octocorallia: Alcyoniidae and Nidalliidae), with descriptions of two new genera. Zoologische
Verhandelingen 245:1-131.

WILLIAMS, G.C. 1992a. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals
(Coelenterata, Anthozoa). Annals of the South African Museum 100(3):249-358.

. 1992b. Revision of the soft coral genus Minabea (Octocorallia: Alcyoniidae) with new taxa from the

Indo-West Pacific. Proceedings of the California Academy of Sciences 48(1):1—26.

. 1996. Octocorallia — Octocorals. Pp. 32-60 in Coral reef animals of the Indo-Pacific—animal life from

Africa to Hawai’i exclusive of the vertebrates, T. M. Gosliner, D. W. Behrens, and G. C. Williams. Sea

Challengers, Monterey. 314 pp.

208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 16

. 2000a. Two new genera of soft corals (Anthozoa: Alcyoniidae) from South Africa, with a discussion of

diversity and endemism in the southern African octocorallian fauna. Proceedings of the California Acad-

emy of Sciences 52(6):65—75.

.2000b. A new species of the soft coral genus Eleutherobia Piitter, 1900 (Coelenterata: Alcyonacea)
from the Tonga Islands. Proceedings of the California Academy of Sciences 52(13):159-169.

WILLIAMS, G. C. AND P. ALDERSLADE. 1999. Revisionary systematics of the western Pacific soft coral genus
Minabea (Octocorallia: Alcyoniidae), with descriptions of a related new genus and species from the
Indo-Pacific. Proceedings of the California Academy of Sciences 51(7):337—364.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 17, pp. 209-225, 10 figs., 1 table. SEP 0 5 200) August 21, 2001

First Record of a Bioluminescent Soft Coral: Description of a
Disjunct Population of Eleutherobia grayi (Thomson and Dean,
1921) from the Solomon Islands, with a Review of
Bioluminescence in the Octocorallia

by

Gary C. Williams
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

A population of alcyoniid soft corals assignable to the species Eleutherobia grayi (Thomson
and Dean, 1931) is described from the Solomon Islands in the southwestern Pacific Ocean,
a taxon previously known only from the Indonesian and Ryukyuan Archipelagos. The spe-
cies is remarkable in that the anthocodiae are strikingly bioluminescent. This represents
the first record of a bioluminescent soft coral, other than a dubious record for Alcyonium.
Other octocorals for which bioluminescence has been verified are restricted to the
gorgonian family Isididae and the Pennatulacea. A table of all octocorallian taxa in which
bioluminescence has been recorded (along with newly presented data) is also included,
along with a brief review of bioluminescence in the Octocorallia. Two additional species of
Eleutherobia, which are presently unidentified and presumably undescribed, one from
Palau, and the other from the Philippine and Mariana Archipelagos, are here reported to
also bioluminesce.

Previously, bioluminescence in octocorals has been presumed to be restricted to many, but not
all, sea pens (Pennatulacea) and two to four species of gorgonians (Alcyonacea: Isididae) (Harvey
1952; Muzik 1978). The present paper reports the discovery of bioluminescence in a species of the
alcyoniid genus Eleutherobia from the Solomon Islands. The discovery represents the first record of
bioluminescence in a soft coral.

The genus Eleutherobia was last revised by Verseveldt and Bayer (1988). Three species of
Eleutherobia have recently been described: E. /utea Benayahu and Schleyer, 1995, from Natal, South
Africa; E. zanahoria Williams, 2000, from the Tonga Islands; and E. vinadigitaria Williams and Lit-
tle, 2001, also from Natal, South Africa. Seventeen described species of the genus (considered valid
taxa) are known from southern A frica and the Indo-West Pacific—four from southern A frica and thir-
teen from the Indo-West Pacific (Somalia to Japan and Tonga). Bioluminescence has not been re-
ported in any previously described species of the genus.

Material collected from a disjunct population of a soft coral species identified as Eleutherobia
grayi (Thomson and Dean, 1931) is described from the Solomon Islands. The specimens do not differ
morphologically from the lectotype (redescribed by Verseveldt and Bayer 1988), which was clearly
illustrated with scanning electron micrographs of sclerites. However, the Solomon Islands material
differs ecologically in one respect from the type material and other known specimens. The previously
known material: type specimens from Indonesia, and a record from the Ryukyu Islands (Verseveldt
and Bayer 1988:33), was collected from sandy or rubbly bottom substrata (sand with small stones and
shells or pieces of dead coral), whereas specimens from the newly discovered population in the Solo-

209

210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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mon Islands have been observed only on hard substrata (floors, walls and ceilings of limestone caves,
alcoves, and overhangs). The populations also differ bathymetrically. The Indonesian and Ryukyuan
populations are known from 30-73 m, while the Solomon Islands population has been observed at
shallower depths, 5—18 m.

Williams (2000:159) has summarized recent discoveries pertaining to natural products biochem-
istry and the genus Eleutherobia.

MATERIAL AND METHODS

Material for this study was collected by SCUBA and preserved directly in 70% ethanol. Sclerites
were isolated by disassociating them from the coenenchyme with household bleach (sodium
hypochlorite). Underwater photographs for Figure | were made with a Nikonis-V camera and Nikonis
SB 103 flash unit. Other photographs and micrographs were made using a Nikon Coolpix 990 digital
camera and a Nikon SMZ-10 dissecting microscope. Scanning electron micrographs were made with
a Leo 1400 series scanning electron microscope. Sclerite drawings were made using an Olympus
CH-2 compound microscope with an attached drawing tube. Digital images and plates of photo-
graphs, micrographs, and scanning electron micrographs were made using Adobe Photoshop soft-
ware. An abbreviation used in the text is CAS (California Academy of Sciences, San Francisco).

SYSTEMATIC ACCOUNT
Family Alcyoniidae Lamouroux, 1812

Eleutherobia Pitter, 1900

Eleutherobia Putter, 1900:449. Verseveldt and Bayer, 1988:27. Williams, 1992:306; 2000:160. Williams and
Little, 2001.

DIAGNOSIS. — Alcyoniid soft corals, colonies digitiform (finger-like), conical or cylindrical,
rarely lobate to subglobular. Polyps monomorphic with calyces absent, however, retracted polyps
may form low rounded or conspicuous and mound-like protuberances of the coenenchyme. Sclerites
predominantly derived from radiates, with spindles, barrels, tuberculate spheroids, rod-like forms, or
crosses sometimes present. Anthocodial sclerites present as crown and points, or as eight points only,
or altogether absent. Color variable.

TYPE SPECIES. — Eleutherobia japonica Pitter, 1900, by monotypy.

DIVERSITY AND DISTRIBUTION. — Seventeen species in the Indo-West Pacific and South Africa
(eastern and southern Africa to Japan, Saipan, and Tonga) (Fig. 10B).

Eleutherobia grayi (Thomson and Dean, 1931)
Figs. 1-10

Nidalia grayi Thomson and Dean, 1931:37, pl. 2 fig. 2. Type locality: Indonesia.
Eleutherobia grayi : Verseveldt and Bayer, 1988:33, figs. 24, 25.
Eleutherobia sp.: Williams, 1996:34 (color photographs taken both at night and during the day).

MATERIAL EXAMINED. — CAS 101096, station number 35, Solomon Islands, Mborokua Island
(Murray’s Island), 18 m depth, 9 November 1994, collected by G. C. Williams with aid of SCUBA,
one whole specimen, 22 mm in length. CAS 147475, same data as CAS 101096, one specimen cut
longitudinally into two halves, 29 mm in length. CAS 147476, station number 16, Solomon Islands,

WILLIAMS: BIOLUMINESCENT SOFT CORAL

FIGURE |. Eleutherobia grayi. A-B. Underwater photographs of living soft corals, Solomon Islands. Colonies (excluding
polyps) are approximately 25 mm in length. A. Photograph taken at night. B. Photograph taken during midday. C. Lectotype,
approximately 33 mm in length (from Thomson and Dean 1931, pl. 2 fig. 2).

Di PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Mborokua Island (Murray’s Island), 10 m depth, 12 September 1998, collected by G. C. Williams
with aid of SCUBA, one whole specimen, 24 mm in length. CAS 101095, station number 20, Solo-
mon Islands, Mborokua Island (Murray’s Island), 16 m depth, 4 November 1994, collected by G. C.
Williams with aid of SCUBA, three whole specimens: 22 mm, 24 mm, and 27 mm in length.

DESCRIPTION. — Growth form and size. Alcyoniid soft corals in which the colonies are
digitiform (Figs. 1A, 2E) or somewhat lobate (Figs. 1B, 2A—D), mostly cylindrical in shape (Fig. 2E),
not markedly tapered, distal apex bluntly rounded (Figs. 1B, 2B—E), somewhat truncate (Figs. 1C,
2A) or somewhat clavate (Fig. 2A, C, D), not pointed. The stalk is very short, as the polyps are distrib-
uted over approximately 95% of surface of colony (Figs. 1, 2). The polyparies of most colonies arise
directly from a broad holdfast (Figs. 1C, 2A). Wet-preserved colonies vary in length from 11 to
29 mm.

Polyps. The polyps are arranged uniformly over the surface of the colonies. They are
monomorphic and do not have calyces, although the retracted polyps may form low rounded or
mound-like protuberances on the surface of the polypary in some preserved or tightly retracted colo-
nies (Fig. 1C). These protuberances are formed from the coenenchyme and hence cannot be defined as
true calyces. In most cases, the retracted polyps are often more or less flush with the surface of the
polypary (Figs. 1B, 2). Polyp sclerites are absent (Figs. 1A, 3, 4).

The living expanded polyps are colorless and translucent, 9-12 mm in length. The length of phar-
ynx is approximately one-third the length of the body of the polyp (Figs. 1A, 3). The peristome is a lus-
trous and reflective opaque white, while the glandular uppermost portions of the mesenterial
filaments are pale yellowish or cream-colored (Fig. 1A). The narrow elongate tentacles are mostly
6-8 mm in length, gradually taper to a point, and have two opposite rows of approximately 12-16
pinnules. (Figs. 1A, 3, 4). Wet-preserved expanded polyps are opaque white (Fig. 2B).

Sclerites. Sclerites are of several distinct types varying from 0.05 to 0.15 mm in length. In the
surface coenenchyme of the polypary and holdfast are seven-radiates (Figs. 5C; 6G; 7E, O; 81),
eight-radiates (Figs. SA, C; 6C, I; 71, K; 8B, F, N), crosses (Figs. 7J, 8D), quadriradiates with three tu-
bercles in one plane and one tubercle vertically disposed in the center (Fig. 6A), and triradiates (Fig.
6K). In the subsurface coenenchyme and deep interior of the colonies are tuberculate rods and irregu-
larly-shaped, somewhat flattened, rod-like forms (Figs. 5B, D; 7G, L, P).

Color. Most colonies are brick red (Fig. 1A), but some have varying amounts of yellow color-
ation (Fig. 1B). Sclerite color varies from red to orange, or colorless.

DISTRIBUTION. — Solomon Islands (5—18 m depth) (present study) plus the type localities:
Ceram Sea, between Misool and the western end of New Guinea; Indonesia (32 m depth) (lectotype,
designated from two syntypes by Verseveldt and Bayer, 1988); and Flores Sea, northwestern end of
Sumbawa, Indonesia (73 m depth) [paralectotype, designated from two syntypes by Verseveldt and
Bayer (1988)]. They also reported the species from the region of Okinawa in the Ryukyu Islands, Ja-
pan (30 m depth). I have examined a large number of specimens of material assignable to the genus
Eleutherobia from Okinawa. These specimens differ in several respects from Eleutherobia grayi, and
are here considered to represent another (as yet undetermined) species of the genus. I therefore con-
sider the occurrence of Eleutherobia grayi in the Ryukyu Archipelago as unverified (Fig. 10).

VARIABILITY. — Colony shape is variable—digitiform, or bilobate, to somewhat globular.
Color is also variable depending on the proportions of red, orange, and colorless sclerites present in
the coenenchyme. Some colonies are a uniform brick red, or red with yellowish polyp mounds, while
others are very pale pink with cream-colored mounds created by the retracted polyps. These ca-
lyx-like mounds are usually uniformly colored, but may be bicolored in some colonies—yellow with
eight radiating red stripes, or red with yellow stripes (Fig. 1).

BIOLUMINESCENCE. — During night dives on three occasions (November 1993, November
1994, and September 1998), a vivid green bioluminescence (bright green flash) was observed imme-
diately upon tactile contact with the fully expanded polyps of Eleutherobia grayi. This light seems to

WILLIAMS: BIOLUMINESCENT SOFT CORAL 213

FIGURE 2. Eleutherobia grayi. A. Wet-preserved specimen (CAS 101096); 22 mm in length. B. Wet-preserved specimen
with expanded polyps (CAS 147476); 24 mm in length. C. Wet-preserved specimen (CAS 147475), external view; 29 mm in
length. D. Wet-preserved specimen (CAS 147475), cut longitudinally to show internal aspects; 29 mm in length. E.
Wet-preserved specimen (CAS 101095); 27 mm in length. Scale bar = 12 mm.

emanate from the region of the peristome (oral disc) and the distal-most region of the pharynx (Figs.
1A, 3, 4), although the precise origin of luminescence remains uncertain. The peristome appears lus-
trous white under the white-light of an underwater camera flash unit, and thus displays a striking con-
trast to other parts of the soft coral colony (Fig. 1A). Research using epiflourescence microscopy to
determine the location of photocytes in this species is in progress.

214

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 17

Pk tet te MA oc > «
FOE EPR eh aha:
: ch SES SUP bre eat :

FIGURE 3. Eleutherobia grayi. A single polyp, fully extended; scale bar = 3 mm.

WILLIAMS: BIOLUMINESCENT SOFT CORAL PRLS

FIGURE 4. Eleutherobia grayi. Mouth and tentacles showing oral disc surrounding the mouth (peristome), which is the sus-
pected region of bioluminescence; scale bar = 5 mm.

Since the polyps of Eleutherobia grayi are bioluminescent and devoid of scleritic armature as
well, it is possible that the attribute of bioluminescence precludes the need for anthocodial armature,
and may be regarded as a defense against potential predators of the soft coral polyps. Only one other
described species of Eleutherobia (E. zanahoria Williams, 2000) is known to lack polyp armature, but
the presence or absence of bioluminescence in this taxon is not known.

ECOLOGICAL OBSERVATIONS. — Much of the shallow water, hard coral cover at the type locality
was dramatically altered between 1994 and 1998, presumably by a combination of warming events
and a series of severe storms. Physical decimation of hermatypic as well as ahermatypic scleractinians
took place, together with a substantial amount of bleaching (predominantly on the distal-most extrem-
ities of the coral colonies). On the exposed reef flats and slopes, replacement of hermatypic
scleractinians by soft corals such as Paralemnalia spp. and various taxa of coralline algae, was ob-

216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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served during this period. In the limestone caves, alcoves, and overhangs (Fig. 9), a physical replace-
ment of Tubastraea faulkneri Wells, 1982 (which provided the dominant cover in 1993 and 1994) by
the soft coral Eleutherobia grayi, was also observed during the same period.

The polyps of the soft coral are retracted into the body of the soft coral during daylight hours (Fig.
1B), gradually expand at dusk, and remain fully expanded (in feeding mode) for the duration of the
night (Fig. 1A).

Several colonies of Eleutherobia grayi were observed to have minute epizoites (mostly tunicates
or bryozoans) growing on the surface of the polyparies, between the polyps.

REMARKS. — Morphologically, material from the Solomon Islands population fully agrees with
the original description of Eleutherobia grayi by Thomson and Dean (1931), and the well-illustrated
redescription of the designated lectotype by Verseveldt and Bayer (1988). Several important points of
concurrence are listed below.

1. Total lack of sclerites in the anthocodiae.

2. Red or yellow coloration due to varying amounts of red and yellow sclerites.

3. Deep interior of colonies with many sclerites, almost exclusively rod-like forms.

4. Size range of sclerites: 0.05—0.15 mm (present study); 0.03—0.18 mm (Verseveldt and Bayer,
1988).

5. Sclerite types: Eight-radiates, seven-radiates, triradiates, crosses, quadriradiates with three tu-
bercles in one plane and one tubercle arising vertically from the center, and irregularly-shaped tuber-
culate rods.

DISCUSSION

BIOLUMINESCENCE IN OCTOCORALS (Table 1). — For the sake of the following discussion, the
various groups of octocorals are defined as follows. Stoloniferous octocorals have separate polyps
connected at their bases by membranous or ribbon-like stolons. Soft corals have polyps embedded in
a common coenenchyme and are attached to hard substrata by basal holdfasts, they have no axial de-
velopment, and only free sclerites comprise the skeletal elements. Gorgonians, like the soft corals,
have polyps embedded in a common coenenchyme and are attached to the substratum by basal hold-
fasts, but unlike soft corals, in addition to free sclerites, have some form of internal axial development
composed of calcium carbonate, gorgonin, or a combination of the two. Sea pens or pennatulaceans
have the coral colony divided into a proximal muscular peduncle that is anchored in soft substrata, and
a distal rachis that contains several kinds of polyps. They may or may not have a calcareous axial skel-
eton.

Williams (1999:23, 49-50) provided a historical review and comprehensive bibliography per-
taining to pennatulacean bioluminescence. The scientific literature regarding bioluminescence in
octocorals is relatively rich, extending back to the sixteenth century with the works of Gesner,
Boussuet, Imperato, Rondelet, and others. Important modern contributions include: Panceri (1871,
1872a, b); Parker (1920); Harvey (1940, 1952); Nicol (1958); Titschak (1965, 1966); Morin (1974,
1976); Muzik (1978); and Herring (1991). Harvey (1952:168) stated, “... but among the Alcyonaria
are to be found some of the most brilliant and striking luminous animals. Of the three groups of
Alcyonaria, the Alcyonacea, the Gorgonacea, and the Pennatulacea, only luminescence of the
Pennatulacea has been carefully studied.” Pertaining to alcyonaceans, only a few species in the
gorgonian family Isididae have been known to luminesce. Mangold’s (1910) record of Leuckart’s lu-
minescent Alcyonium is considered by Harvey (1952:169—170) to be “very dubious.” No subsequent
records of bioluminescence in the genus A/cyonium, or any other soft coral taxon for that matter, are
known in the previous literature.

The only records of bioluminescent soft corals are represented by new data presented here of
Eleutherobia grayi from the Solomon Islands, as well as observations made on two other species of

WILLIAMS: BIOLUMINESCENT SOFT CORAL

TABLE |. Records of bioluminescence in octocorals.

Taxon

Order Alcyonacea
Soft corals
ALCYONIIDAE
Eleutherobia grayi
Eleutherobia sp. indet. #1
(Luzon and Saipan)
Eleutherobia sp. indet. #2
(Palau)

Gorgonians
ISIDIDAE
Lepidisis olapa
Isidella elongata
Keratoisis sp.
Primnoisis sp.

Order Pennatulacea
Sea Pens
VERETILLIDAE
Cavernularia habereri
Cavernularia pusilla
Veretillum cf. manillense
Veretillum cynomorium
ECHINOPTILIDAE
Actinoptilum molle
RENILLIDAE
Renilla muelleri
Renilla koellikeri
Renilla reniformis
FUNICULINIDAE
Funiculina quadrangularis
PROTOPTILIDAE
Distichoptilum gracile
UMBELLULIDAE
Umbellula huxleyi
Umbellula thomsoni
VIRGULARIDAE
Stylatula elongata
Acanthoptilum gracile
Virgularia mirabilis
PENNATULIDAE
Ptilosarcus gurneyi
Pennatula phosphorea
Pennatula rubra
Pteroeides spinosum

Depth

60-2600 m

650-4300 m
210-6100 m

9-50 m
10-146 m
9-400 m

0-68 m
300-3609 m

217

Color of light

References on bioluminescence

present study

Gosliner and Starmer - pers. commun.;
Williams - pers. observ.

Starmer - pers. commun.

Muzik (1978:735; 1981[82]:56)
Muzik (1978:735)
Harvey (1952:169)
Harvey (1952:169)

Harvey (1917)

Panceri (1872a)

Williams - pers. observ.

Bujor (1901); Titschak (1965)

Williams (1990:63)

Parker (1920)
Morin (1976:632)
Ward and Cormier (1978)

Thomson (1874:149); Herring (1991)
Herring (1991)

Herring (1991)
Tizard et al. (1885:49)

Morin (1976:632)
Morin (1976:630)
Herdman (1913); Nicol (1958)

Morin (1976:630)

Herdman (1913); Titschak (1966)
Panceri (1871); Titschak (1966)
Panceri (1871); Titschak (1966)

218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 5. Eleutherobia grayi sp. nov. Sclerites. A. Polypary surface. B. Polypary interior. C. Stalk surface. D. Stalk interior.
Scale bar = 0.1 mm.

WILLIAMS: BIOLUMINESCENT SOFT CORAL a)

FIGURE 6. Eleutherobia grayi sp. nov. Scanning electron micrographs of coenenchymal sclerites from the mound-like protu-
berances formed by the retracted polyps. A. 0.04 mm. B. 0.05 mm. C. 0.06 mm. D. 0.05 mm. E. 0.07 mm. F. 0.06 mm. G.
0.07 mm. H. 0.07 mm. I. 0.07 mm. J. 0.07 mm. K. 0.04 mm.

Eleutherobia (Eleutherobia spp. indet.). One of these is found on vertical surfaces at 12 m depth in
southern Luzon, Philippines (pers. observ., and pers. commun. T. M. Gosliner), and Saipan (pers.
commun. J. Starmer), and the other has been collected from Palau (pers. commun. J. Starmer). These
observations presented here are the first records of bioluminescent soft corals (outside of the dubious
record of Leuckart’s Alcyonium).

The phenomenon of bioluminescence in the Pennatulacea, although commonly encountered in
bathymetrically diverse habitats and supported by a relatively rich literature, is by no means universal,

220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGURE 7. Eleutherobia grayi sp. nov. Scanning electron micrographs of coenenchymal sclerites from the surface and
subsurface of the polypary. A. 0.09 mm. B. 0.08 mm. C. 0.05 mm. D. 0.08 mm. E. 0.09 mm. F. 0.08 mm. G. 0.08 mm. H.
0.07 mm. I. 0.06 mm. J. 0.10 mm. K. 0.08 mm. L. 0.09 mm. M. 0.05 mm. N. 0.05 mm. O. 0.08 mm. P. 0.07 mm.

WILLIAMS: BIOLUMINESCENT SOFT CORAL

FiGureE 8. Eleutherobia grayi. Scanning electron micrographs of coenenchymal sclerites from the surface of the holdfast re-
gion of the stalk. A. 0.09 mm. B. 0.08 mm. C. 0.07 mm. D. 0.08 mm. E. 0.08 mm. F. 0.08 mm. G. 0.05 mm. H. 0.08 mm. I.
0.10 mm. J. 0.11 mm. K. 0.10 mm. L. 0.07 mm. M. 0.13 mm. N. 0.11 mm.

222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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limestone wall

cave or alcove

colonies of

Eleutherobia grayi

Depth (m)

interior of
sandy bottom limestone wall

11.0

FIGURE 9. Habitat Section. Diagram of longitudinal section through a limestone wall at approximately 10 meters in depth,
Mborokua Island, Solomon Islands, showing disposition of densely-set colonies of Eleutherobia grayi. Depths shown at left
are in meters.

as some taxa are definitely not luminescent. For example, Herring (1978:204) stated that some species
of Virgularia have been shown to be non-luminous.

Bioluminescence has thus far been recorded in only 53% of the pennatulacean families (8 of 15),
40% of the genera (13 of 32), and 10% of the estimated number of valid species (19 of 186). If the total
number of described species in the literature are taken into account, then the latter percentage drops to
approximately 4% (19 of 436). These estimates represent minimum values, but could be much higher,
and are based on numbers of taxa in Williams (1995:93) and Table 1 of the present paper.

[t is surprising that after nearly 450 years of published observations resulting in more than eighty
published accounts of pennatulacean bioluminescence, a mere nineteen species (or approximately
10% of the valid species) have been recorded. This number seems especially low considering the in-
troduction of modern technological means to make observations—such as SCUBA, remote opera-
tional vehicles (ROV’s), and manned deep-sea submersibles. Only a few sea pens are diurnal and/or

WILLIAMS: BIOLUMINESCENT SOFT CORAL 223

Bougainville

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oe

wf PACIFIC OCEAN
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Shortland :

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Santa Isabel

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- 4¢ -
New Georgia Group Bios 2 Malaita
Mborokua

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FiGuRE 10. A. Map of the Solomon Archipelago, southwestern Pacific Ocean; dotted line marks the political boundary be-
tween Papua New Guinea (upper left) and the Solomon Islands. B. Map of the Indo-West Pacific showing geographic distribu-
tion of the genus Eleutherobia;, @ = collecting stations; arrows show collecting stations for Eleutherobia grayi.

224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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zooxanthellate, and are not known to bioluminesce. The majority of shallow water species (at least in
the Indo-Pacific) are nocturnal and azooxanthellate (Williams, pers. observ.). One South African spe-
cies, Actinoptilum molle, appears to be active both during the day and night, and is at the same time
azooxanthellate and bioluminescent (Williams, 1990:63). In addition, a remarkable diversity of
deep-water taxa are known (Williams 1993:733—734; 1997:499, 503) that live in perpetual darkness.
It is therefore assumed that the actual number of bioluminescent sea pens could be much higher than is
presently documented.

ACKNOWLEDGMENTS

I am grateful to James Kasson (Menlo Park, California), John Starmer (Koror, Palau), and
Terrence M. Gosliner, Alan Dekelboum, and Roy Eisenhardt (California Academy of Sciences) for
their underwater observations. I thank James Morin (Cornell University) and Stephen Cairns (Smith-
sonian Institution) for their insight and useful comments.

LITERATURE CITED

BENAYAHU, Y. AND M. H. SCHLEYER. 1995. Corals of the south-west Indian Ocean II. E/eutherobia aurea spec.
nov. (Cnidaria, Alcyonaria) from deep reefs on the KwaZulu-Natal Coast, South Africa. South African As-
sociation for Marine Biological Research, Oceanographic Research Institute, Investigational Report
68:1-12.

BuJor, P. 1901. Sur organisation de la Vérétille. Archives de zoologie expérimentale et générale, notes et revur
(Ser. 3) 9: No. 4, xlix—Ix.

HARVEY, E. N. 1917. Studies on bioluminescence. 6. Light production by a Japanese pennatulid Cavernularia
habereri. American Journal of Physiology 42:349-358.

. 1940. Living Light. Princeton University Press, Princeton, New Jersey. 328 pp.

. 1952. Bioluminescence. Academic Press, New York. 649 pp.

HERDMAN, W. A. 1913. “Phosphorescence” of Pennatulida. Nature, London 91:582.

HERRING, P. J., ed. 1978. Bioluminescence in action. Academic Press, London. 570 pp.

. 1991. Observations on bioluminescence in some deep-water anthozoans. Hydrobiologia
216/217:573-579.

MANGOLD, E. 1910. Die Produktion von Licht. Pp. 225—392 in Handbuch der vergleichende Physiologie, Jena 3
(2nd half), H. Winterstein, ed.

Morin, J. G. 1974. Coelenterate bioluminescence. /n Coelenterate Biology: Reviews and New Perspectives, L.
Muscatine and H. Lenhoff, eds. Academic Press, New York. 501 pp.

. 1976. Probable functions of biolumninescence in the Pennatulacea (Cnidaria, Anthozoa). /n
Coelenterate ecology and behavior, G.O. Mackie, ed. Plenum, New York. 744 pp.

MUZIK, K. 1978. A bioluminescent gorgonian, Lepidisis olapa, new species (Coelenterata: Octocorallia), from
Hawaii. Bulletin of Marine Science 28(4):735—741.

. 1981(82). Proceedings of the Fourth International Coral Reef Symposium 2:756 [abstract].

NICOL, J. A. C. 1958. Observations on the luminescence of Pennatula phosphorea, with a note on the lumines-
cence of Virgularia mirabilis. Journal of the Marine Biological Association of the United Kingdom
37:55 1-563.

PANCERI, P. 1871. Gli organi luminoie la luce delle Pennatule. Rendiconto dell’ Accademia della scienze, Napoli
10(1):204-211.

. 1872a. Etudes sur la Phosphorescence des Animaux Marins. II. Du siége du mouvement lumineux dans
les Méduses; III. Organes lumineux et lumiére des Pennatules; VI. Sur un Pennatulaire phosphoresenct en-
core inconnu dans les environs de Naples (Cavernularia pusilla); 1X. Des organes lumineux et de la lumiére
des Béroidiens. Annales des Sciences Naturelles, sér. 5 (Zoologie)16 (8): 1-66.

————. 1872b. The luminous organs and light of the Pennatulae. Quarterly Journal of Microscopical Science,
London 12:248-254. [English translation of Panceri, 1871]

PARKER, G. H. 1920. The phosphorescence of Renilla. Proceedings of the American Philosophical Society
19:171-175.

WILLIAMS: BIOLUMINESCENT SOFT CORAL 225

THOMSON, C. W. 1878. 1874. The depths of the sea—an account of the general results of the dredging cruises of
H.M.S. “Porcupine” and “Lightning” during the summers of 1868, 1869, and 1870, under the scientific di-
rection of Dr. Carpenter, F.R.S., J. Gwyn Jeffreys, F.R.S., and Dr. Wyville Thomson, F.R.S., 2nd ed.
Macmillan and Company, London. 527 pp.

THOMSON, J. A. AND L. M. I. DEAN. 1931. The Alcyonacea of the Siboga Expedition with an addendum to the
Gorgonacea. Siboga Expedition Monographs 13d:1—227.

TITSCHAK, H. 1965. Untersuchungen tuber das Leuchten der Seefeder Veretillum cynomorium (Pallas). Vie et Mi-
lieu 15:547-563.

. 1966. Uber die Lumineszenz und ihre Lokalisation bei Seefedern. Zoologischer Anzeiger,
Supplementband 29, 1965(1966):120-131.

TIZARD, T. H., H. N. MOSELEY, H. Y. BUCHANAN, AND J. MARRAY. 1885. Narrative of the cruise of H.M.S. Chal-
lenger with a general account of the scientific results of the expedition. Report on the Scientific Results of
the Voyage of the H.M.S. Challenger during the years 1873-76, Narrative, Vol. | (first part):1—509.

VERSEVELDT, J. AND F. M. BAYER. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and
Nidaliopsis (Octocorallia: Alcyoniidae and Nidalliidae), with descriptions of two new genera. Zoologische
Verhandelingen 245:1—131.

WARD, W.W. AND M. J. CORMIER. 1978. Energy transfer via protein protein interaction in Renilla
bioluminescence. Photochemistry and Photobiology 27(4):389-396.

WILLIAMS, G. C. 1990. The Pennatulacea of southern A frica (Coelenterata, Anthozoa). Annals of the South A fri-
can Museum 99(4):31—119.

. 1992. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals (Coelenterata,

Anthozoa). Annals of the South African Museum 100(3):249-358.

. 1993. Biotic diversity, biogeography, and phylogeny of pennatulacean octocorals associated with coral

reefs in the Indo-Pacific. Proceedings of the Seventh International Coral Reef Symposium 2:729-735.

. 1995. Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and synop-

ses. Zoological Journal of the Linnean Society, London 113:93—140.

. 1996. Octocorallia—Octocorals. /n Coral reef animals of the Indo-Pacific, animal life from Africa to

Hawai’i exclusive of the vertebrates, T. M. Gosliner, D. W. Behrens, and G. C. Williams. Sea Challengers,

Monterey, California. 314 pp.

. 1997. Preliminary assessment of the phylogeny of Pennatulacaea (Anthozoa: Octocorallia), with a

reevalutation of Ediacaran frond-like fossils, and a synopsis of the history of evolutionary thought regarding

the sea pens. Proceedings of the Sixth International Conference on Coelenterate Biology: 497-509.

. 1999. Index Pennatulacea—annotated bibliography and indexes of the sea pens (Coelenterata:

Octocorallia) of the world 1469-1999. Proceedings of the California Academy of Sciences 51(2):19-103.

. 2000. A new species of the soft coral genus Eleutherobia Piitter, 1900 (Coelenterata: Alcyoniidae) from
the Tonga Islands. Proceedings of the California Academy of Sciences 52 (13):159-169.

WILLIAMS, G. C. AND S. A. LITTLE. 2001. A new species of the soft coral genus Eleutherobia Piitter, 1900
(Octocorallia: Alcyoniidae) from South Africa. Proceedings of the California Academy of Sciences
52(16):193—206.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 18, pp. 227-244, 11 figs., 1 table. October 26, 2001

Four New Species of Forcepia (Porifera, Demospongiae,
Poecilosclerida, Coelosphaeridae) from California, and Synonymy —__
of Wilsa de Laubenfels, 1930, with Forcepia, Carter, 1874°)0° °°.

Welton L. Lee
Fellow, California Academy of Sciences | oe
Golden Gate Park, San Francisco, California 94118 _ or eM 02543

Four new species of the marine sponge genus Forcepia (Porifera, Demospongiae,
Poecilosclerida, Myxillina, Coelosphaeridae), are described and Wilsa hymena de
Laubenfels, 1930, which is now designated as Forcepia (Forcepia) hymena (de Laubenfels,
1930), is redescribed on the basis of new material found in the collections of the Scripps In-
stitution of Oceanography. As a result of detailed comparison of this material with de
Laubenfels’ type, Wilsa is synonymized with Forcepia. Heretofore, only one species of this
group (Wilsa hymena) had been described from California. Full descriptions, figures and
relevant morphological and habitat information have been included.

by DEC 0.3 2007

While the rich coastal waters along the California coast support a vast and varied invertebrate
fauna, our knowledge of the organisms found in this region remains disappointingly poor. A prime ex-
ample is the marine sponge fauna. Those studying sponges must rely largely on a publication from
1932 by de Laubenfels and a field manual on intertidal invertebrates last published in 1975 by
Hartman as their main source of information. Scattered papers, of which only a few are recent, provide
the remainder of the available information.

Five years ago, the David and Lucile Packard Foundation generously funded the author for a
study of the sponges of California. That study has been completed (Lee, Elvin, Reiswig, in prepara-
tion). Over 250 species have been found in Californian waters, many of these new to science. The
present paper represents the first of an anticipated series describing the newly discovered species.

In the past, sponges bearing forceps as microscleres have been variously placed in different taxa.
From recent investigations by Van Soest (pers. commun.) and Hajdu and Vacelet (pers. commun.)
forceps-bearing taxa are now largely relegated to the genus Forcepia Carter, 1874 (family Coelo-
sphaeridae). An exception is the genus Asbestopluma, Norman, 1882 (family Cladorhizidae), which
consists largely of abyssal sponges with forceps of different structure, not considered homologous
with the forceps of Forcepia. The papers by the above mentioned authors are part of a worldwide ef-
fort to review and revise all presently used taxonomic categories through a re-examination of the type
material on which these taxa are based. The papers will be compiled in the Systema Porifera to be pub-
lished this year (Hooper and Van Soest, in press). The revision of Forcepia includes two newly de-
fined subgenera, Forcepia and Leptolabis, the latter distinguished by the presence of basal
acanthostyles and a hymedesmoid-like skeletal architecture. The California sponges described here
all belong to the subgenus, Forcepia.

To date only two forceps-bearing sponges have been reported from California, Asbestopluma
lycopodium (Levinsen, 1886) and Wilsa hymena de Laubenfels, 1930. The first is a member of the
family Cladorhizidae, and will not be discussed here. The genus Wil/sa was erected by de Laubenfels

Zo]

228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

in 1930 for a single forceps-bearing specimen. Recent review of newly found material suggests that
this species belongs to the genus Forcepia and de Laubenfels’ Wilsa is synonymized with Forcepia.
Besides aredescription of Wilsa hymena, four new species of Forcepia are described in this paper.

MATERIALS AND METHODS

Material examined in this study was predominantly taken from museum collections. Abbrevia-
tions for specimens in these collections are as follows: NHM, The Natural History Museum, London;
CASIZ, California Academy of Sciences; BIC-SIO, Scripps Institution of Oceanography, Inverte-
brate Collection; SBMNH, Santa Barbara Natural History Museum; U.S.N.M., U.S. National Mu-
seum, Smithsonian Institution; YPM, Peabody Museum, Yale University.

Spicule preparations and cross-sections were routinely made according to the procedures of
Hartman (1975). Slide preparations were mounted in Permount. All measurements, including
spicules, were made with a stage micrometer directly through a compound microscope. Width mea-
surements for megascleres were taken at the thickest point of the spicule shaft. Isochela lengths were
taken from the apices of alae; length measurements of other spicules refer to maximum lengths. No
less than 50 measurements were made for each spicule type and the data subjected to statistical analy-
ses of range and mean. These measurements were displayed graphically to determine if distinct
spicule size classes were present. Distinctive size classes are deemed legitimate only when these
graphs show either non-overlapping, or distinct bi- or tri-modal distributions.

Spicule measurements are shown in this paper with the lowest size listed first and the greatest size
listed last. In cases where only one specimen was measured, the mean is given in between these two
extremes and underlined (234-268-295 um). If more than a single specimen is measured then the
range of means [from lowest to highest] is given and underlined (234—259—271—295 ym).

Spicules were prepared for the scanning electron microscope (SEM) as described above but
mounted and dried on 1.5 cm round slides. These were mounted on stubs with double-sided tape, sput-
ter coated with gold-paladium, and examined on an Hitachi S-520 scanning electron microscope.

SPECIES DESCRIPTIONS

Family Coelosphaeridae Hentschel, 1923

Genus Forcepia Carter, 1874
Wilsa de Laubenfels, 1930:27

Forcepia (Forcepia) acanthostylosa sp. nov.
Figs. | and 2

MATERIAL. — Holotype: SBMNH 345543, U.S.A. California, San Miguel Island, Cuyler Har-
bor, Depth 10.7 to 12.2 m, Collectors: B. Scronce, M. Conboy, C. Carreon, and L. Bray, 19 February
1964.G. E. and N. Macginitie Port Hueneme Collection. Paratype: CASIZ 154368, U.S.A., Califor-
nia, Santa Barbara County, Santa Cruz Island, small cove midway between Chinese Harbor and Pris-
oners Harbor. Depth 10.3 to 12.2 m. Collectors: B. Scronce, M. Conboy, and L. Bray, 3 July 1963.
G. E. and N. Macginitie Port Hueneme Collection.

DISTRIBUTION. — Known only from two localities in southern California: San Miguel Island
(holotype) and Santa Cruz Island (paratype).

HABITAT. — Habitat descriptions were not included in the collection data.

SHAPE. — Holotype, thickly encrusting, |.5—3.3 cm thick. Sponge irregular in shape, 6 cm long
by 2.0-3.8 cm wide. The sponge appears to have encrusted a mat of bottom material, including algae,
other invertebrates and sand. Paratype, thinly encrusting on a shell of the bivalve Hinnites

LEE: CALIFORNIA FORCEPIA (PORIFERA)

FIGURE |. Scanning electron micrographs of the spicules of Forcepia (Forcepia) acanthostylosa, sp. nov. Holotype
(SBMNH 345543) a. Forceps 4,000x, b. Forceps 7,000x, c. Large sigma 2,000x, d. Small sigma 3,000x, e. Small arcuate
isochela 7,000x, f. Large arcuate isochela 4,000, g. Substylote 500~, h. Small acanthostyle 1500*, 1. Large acanthostyle 500~.

FIGURE 2. Forcepia (Forcepia) acanthostylosa, sp. nov. a. Paratype (CASIZ 154368), on fragments of Hinnites
multirugosus 1.0—1.5 mm thick. b. Holotype (SBMNH 345543), dimensions. 1.5—3.25 cm thick, 6.0 x 2.0—3.75 cm wide.

230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

multirugosus, which had been broken into several pieces. The sponge thickness was up to 1.5 mmina
few places, but mostly 1.0 mm or less.

COLOR. — Live—yellow to orange; preserved in ethanol—light tan.

OSCULA. — On surface, randomly distributed on the holotype. Only one seen on the largest piece
of the paratype. Oscula oval or nearly so, with no rims, but appear sunken below the surface. These
range from 0.5 to 1.5 mm in diameter. A few large surface openings are also present. These are some-
what oblong and roughly 2.0 by 4.0 mm; they are probably not oscula.

TEXTURE AND SURFACE CHARACTERISTICS. — Texture firm, but soft, elastic to somewhat com-
pressible. Surface smooth, opaque, somewhat irregularly lobate to verrucose. These surface features
never of a high profile and prominent but usually of low profile and gently undulating.

ECTOSOME. — Made up of parallel tylotes to subtylotes very closely and tightly packed making
the ectosome exceedingly firm. The thickness of this layer ranges from 182 to 200 um. The strength of
the ectosome almost always leads to the tearing of cross-sections.

CHOANOSOME. — Difficult to assess since the firm ectosome tends to drastically disrupt the
choanosome in any cross-sections. Furthermore, presence of dirt, algae and other extraneous material
(less noticeable in the smaller paratype) makes sectioning difficult. What is apparent in the
choanosome is a very loose reticulation of acanthostyles of two size classes which often changes to a
disorganized dispersal of the same. In many cases the choanosome becomes very thick, heavy and dif-
ficult to characterize. In the few relatively open spaces that do exist there is a plethora of microscleres
of all kinds, with sigmas dominating in both numbers and size. Superimposed on the reticulation or
disorganized dispersal of acanthostyles within the choanosome one can often find thick tracts varying
from 46 to 72 um, consisting mostly of acanthostyles of both size classes and, sometimes, tylostyles to
styles. The smaller acanthostyles tend to be either echinating these tracts or involved in the formation
of the reticulation. These tracts appear thickest near the ectosome and dominate the subectosomal
area.

The paratype is relatively free of extraneous material. The holotype is completely invaded by dirt,
algae and other material, making the overall structure difficult to determine. On occasion, areas of
some cross-sections show a basal layer of spongin in which acanthostyles seem to have their heads im-
bedded but this is not at all clear.

MEGASCLERES. — Acanthostyles of two size classes. Small: 68—77—84-101 pm. x
2.4-3.9-5.8-7.3 um (Fig. lh). Spicule straight. Head tends to be flattened and covered with many
spines. Many spines also on the upper third of the shaft. However, some spines occur almost to the tip.
Spines large and robust. With the exception of those spines on the head, almost all are slightly re-
curved toward the head end. Tip spineless, sharply angled and pointed. Large: 181—220—236-265
um. x 2.4-4.4-7.0-9.7 um (Fig li). Spicules straight or with upper third slightly curved. Heads often
flattened. Most spines are on the head and upper fifth of the spicule, a few occurring to near the tip.
Spines mostly small, erect and sharply pointed. Those on the head tend to be crowded and may be
blunt or irregular in shape. Tip free of spines, long and gently angled to a sharp point. Tylotes to
subtylotes of a single size class. 195—227—233-—258 yum. x 3.6—5.0—5.6—6.0 um (Fig. 1g). Spicules
straight, heads usually smoothly rounded but sometimes somewhat elongated. Frequently the shaft is
gently tapered to one end, giving rise to a spicule with unequal ends, one smaller than the other. Some-
times one end is stylote while the other is subtylote.

MICROSCLERES. — Arcuate isochelae of two distinct size classes. Small: 22—26-29-36 um
(Fig. le). Shaft thick and strongly curved. Alae small with edges gently rounded. The lateral alae at-
tached to the shaft most of their length. Large: 46—51—58—68 pm (Fig. If). Shaft thick and strongly
curved. Alae somewhat elongate with edges either nearly square (lateral alae) or somewhat pointed
(frontal alae). Lateral alae clearly detached from the shaft for at least half their length.

Sigmas of two distinct size classes. Both occur in S and C configurations. Small: 26-38-47 um

id). More or less even to slightly asymmetric curvature forming a medium arch. Tips thin,

LEE: CALIFORNIA FORCEPIA (PORIFERA) 231

sharply pointed and angled inwards. One end often twisted out of the plane of the other. Large:
5368-72-78 um (Fig. Ic). More or less even to slightly asymmetric. Much shallower arch. Tips thin,
sharply pointed and angled inwards.

Forceps of two distinct size classes. Small: 7—10—11—13 um (Fig. 1b). Legs not parallel but an-
gled to about 30° from the median between the two equally long legs. Covered with spines with those
at the tip of the legs largest and strongly recurved. Spines on the inner edges of the legs more strongly
recurved than those on the outer edges. Large: 21—24—29 um (Fig. 1a). Legs not parallel, slightly an-
gled to about 10° from the median between the two equally long legs. Covered with numerous small,
angled, erect spines, looking like the teeth of a saw. Those on the inner edge of the legs tend to be
larger than those on the outer edges. The tips of the legs have caps delineated by a ring of small spines.

ETYMOLOGY. — The species is named acanthostylosa to recognize that it is the first Forcepia
from California with acanthostyles.

REMARKS. — This species is the only Forcepia species with acanthostyles reported from the
west coast of North America from Baja California to Canada. Only one Forcepia species has been
noted from this region (Austin and Ott 1987). These authors describe a species similar to Forcepia
(Forcepia) japonica Koltun, 1959, which has styles, but not acanthostyles. However the Canadian
species was noted as having styles to acanthostyles with few spines. This species has only single size
classes of all microsclere types and differs in almost all other respects.

Van Soest (pers. commun.) proposes two subgenera for this genus: Leptolabis for species with a
hymedesmoid kind of structure, with the acanthostyles embedded in a basal layer of spongin, and
Forcepia for those in which the styles or acanthostyles are structural megascleres making up the
choanosomal reticulation. This new species appears to have most of its megascleres involved as struc-
tural elements in a reticulation. It has a thick choanosome showing a reticulate pattern and the obser-
vations of acanthostyles possibly embedded in a basal layer of spongin are too inconclusive to allow
transfer to the subgenus Leptolabis.

Forcepia (Forcepia) elvini sp. nov.
Figs. 3 and 4

MATERIAL. — Holotype: CASIZ 108399, U.S.A., California, Marin Co., Cordell Bank, approx-
imately 20 miles due west of Pt. Reyes. Depth 82.3 m, Collectors: Swift, Smith, Hanna, September
1940.

DISTRIBUTION. — To date only known from the type locality, Central California, Marin Co.,
Cordell Bank.

HABITAT. — Habitat information was not included in the collection data.

SHAPE. -—— Sponge, thick, encrusting, irregular but somewhat rounded; 3.7 cm at the widest
point, 3.5 cm at the narrowest point; 1.5 to 2.5 cm high.

COLOR. — Color in life not recorded; cream white in ethanol.

OSCULA. — Difficult to interpret. Openings, |—3 mm in diameter, round to nearly so with irregu-
lar distribution, abundant at, and flush with the surface. These appear to penetrate well into the interior
of the sponge where smaller, round openings may be seen.

TEXTURE AND SURFACE CHARACTERISTICS. — Texture firm, slightly compressible. Surface su-
~ perficially smooth, some areas with irregularly shaped lobes and others which appear layered with
thin, flat plates. The edges of the lobes and plates are distinctly hispid; the general surface is likewise
hispid, but to a lesser degree.

ECTOSOME. — (Fig. 4a). The ectosome consists of a very thin layer of tightly packed subtylotes
from 24 to 36 um thick; occasionally to 48 um thick. Superimposed over the subtylotes 1s a layer ap-
proximately 24 um thick, packed with microscleres of which isochelae of both size classes appear to
dominate. The presence of the layer of isochelae and the smaller size of the subtylote as compared to

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

FIGURE 3. Scanning electron micrographs of the spicules of Forcepia (Forcepia) elvini sp.nov. Holotype (CASIZ 108399) a.

Large forceps 2,000x, b. Small forceps 2,000x, c. Sigma 1500x, d. Small isochela 4,000x, e. Large isochela 2,000x, f.
Substylote 500, g. Style with spine 300~, h. Style 300~.

‘epia (Forcepia) elvini sp. nov. a. Cross-section 40x. b. Holotype (CASIZ 108399), dimensions 3.5 x 3.7 cm x

LEE: CALIFORNIA FORCEPIA (PORIFERA) 233

the styles in the choanosome make it difficult to see the subtylotes, such that one may initially assume
that the ectosome is made up of microscleres only. The tips of the thick tracts of styles in the
choanosome frequently push through the ectosome and form brushes on the surface.

CHOANOSOME. — (Fig. 4a). The choanosome is formed of a robust reticulation of styles. The re-
ticulation is dominated by thick tracts, normally ranging from 121 to 133 wm but sometimes as thick
as 182 um. Superimposed over these tracts and connecting them is a reticulation of smaller tracts, 61
to 91 um thick. The nodes of this reticulation are particularly thick and noticeable.

MEGASCLERES. — Styles of a single size class. 257-338-393 um x 15—16—18 um (Fig. 3g, h).
Spicules almost always smooth, a few with a very small spine; almost always gently curved near the
center. Shaft thick, slightly thicker near the center. Heads gently rounded, a few may approach a
subtylostyle configuration. Point somewhat sharp.

Subtylotes of a single size class (Fig. 3f). 222-251-335 um x 6—7-10 um. Spicule smooth,
straight to slightly undulate; shaft moderately thick, often slightly wider at one end. Heads very
slightly inflated and smoothly rounded.

MICROSCLERES. — Arcuate isochelae of two size classes. Small: 16—19—21 ym (Fig. 3d). Shaft
thin with well rounded, gentle arch. Alae somewhat smoothly pointed and well separated. Lateral alae
directed rather sharply back towards the shaft; two thirds of their length is attached to the shaft. Large:
26-4449 um (Fig 3e). Shaft thick and strongly arched. Central ala somewhat narrowed with rounded
but even more narrowed tip. Lateral alae wider and well rounded but short relative to length of shaft.
One half of their length is attached to the shaft.

Sigmas of one size class (Fig. 3c). 39-49-56 um. Arch shallow, mostly eccentric with one end
rounded, the other not. Tips sharp; the tip of the rounded end slightly bent inward, the tip on the oppo-
site end sharply bent inward.

Forceps of two size classes. Small: | |—15—28 um (Fig. 3b). Legs often unequal in length, notice-
ably thickest where they join; not parallel but angled to about 30° from the median between the two
legs. Surface not spined but gently undulate. Small caps at the ends of the legs are but slightly inflated
bulbs. Large: 36-49-55 um (Fig. 3a). Legs equal to subequal, very thin except for where they join;
nearly parallel, angled to about 10° to 12° from the median between the two legs. Both interior and ex-
terior surfaces covered with spines, these pointed away from the tip of the foot. Spines sharpest and
most numerous near tip of feet. Distinct saucer shaped caps present.

ETYMOLOGY. — This species is named after Dr. David Elvin, a sponge biologist, computer spe-
cialist, and a long time friend and colleague.

REMARKS. — This species appears to be unique, especially in regard to its two size classes of for-
ceps. The larger appears very similar to those frequently seen in other species, showing numerous
teeth on the legs on both exterior and interior surfaces. The smaller size class is quite different in that it
has an undulating surface with no apparent spines at all. Noteworthy also, is the presence of tiny spines
on some of the styles making up the distinct choanosomal reticulation.

Forcepia (Forcepia) macrostylosa sp. nov.
Figs. 5 and 6

MATERIAL. — Holotype: CASIZ 146074, U.S.A., California MET Sta. 105. Catalina Basin,
33°10/N, 118°36’W, 1271-1280 m, 25’ otter trawl. January 29, 1981, 2400-0230. R/V New Horizon.
Coll. K. Smith, S. Luke.

DISTRIBUTION. — This species is presently known only from its type locality, Southern Califor-
nia, Catalina Basin, California.

HABITAT. — Habitat information was not included in the collection data.

SHAPE. — Thick, massive, somewhat domed-bulbous, 5.0 by 3.5 cm and height to 23 mm.

COLOR. — Color in life unknown; light tan in ethanol.

234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

FIGURE 5. Scanning electron micrographs of the spicules of Forcepia (Forcepia) macrostylosa sp. nov. Holotype (CASIZ
146074) a. Forceps 1500x, b. Sigma 1500x, c. Small arcuate isochela 4000x, d. Large arcuate isochelae 2500x, e. Legs of
forcep 5000x, f. Substylote 300x, g. Subtylostyle 300x, h. Style 300x.

FIGURE 6. Forcepia macrostylosa sp. nov. Holotype (CASIZ 146074) a. Cross-section of the ectosome 40x. Note that the
bundles of subtylotes are worn away except at the extreme right of the photograph. b. Cross-section of the choanosome, 80x.
Note the ladderlike skeleton. c. Holotype, dimensions 5.0 x 3.5 cm, 23 mm high.

LEE: CALIFORNIA FORCEPIA (PORIFERA) 235

OSCULA. — Difficult to discern. May be situated below the surface where the platelike surface
creates openings ranging from 3.0 to 12.5 mm or greater.

TEXTURE AND SURFACE CHARACTERISTICS. — Superficially smooth; some areas totally
smooth, others layered with overlapping thin, flat plates. Consistency spongy, compressible.

ECTOSOME. — (Fig. 6a) Formed of a thin layer of packed subtylotes which is easily removable.
The thickness is usually 48 to 60 um, but sometimes reaches 72 um. In places the ectosomal layer of
subtylotes is worn away.

CHOANOSOME. — (Fig. 6b) The choanosome consists of a ladderlike reticulate skeleton with
tracts of subtylostyles about 91 um. thick. These tracts support a loose reticulation of thinner bundles
ranging up to 24 um thick of from 2 to 5 or more subtylostyles. The thick tracts and ladderlike reticula-
tion becomes less obvious as the ectosome is approached. Near the surface, tracts may become bent al-
most parallel to the surface and the reticulation gets more confused, complicated and random.

MEGASCLERES. — Subtylostyles to styles of a single size class (Fig 5g, h). 268-499-593 um x
18-20-22 um. Smooth, most curved close to the head end. Shaft thick, of more or less even width ex-
cept slightly wider just below the head. Tip hastate to tornote-like, often with a slight expansion of the
shaft before narrowing to the tip.

Subtylotes of a single size class. 309-372-540 um x 6-8-9 um (Fig. Sf). Shaft thin, either
straight or somewhat sinuous with heads abruptly and slightly expanded. Heads often unequal in size,
the smaller set off by a slight constriction of the shaft.

MICROSCLERES. — Arcuate isochelae of two size classes. Small: 19-23-29 um (Fig. 5c). Shaft
with slight or moderate curvature. Alae only somewhat separated. Central ala is the longest; often an-
gled or pointed. Lateral alae smaller, rounded and most often with an obvious, very small, rounded in-
cipient ala next to the shaft. Large: 36—44—S0 um (Fig. 5d). Sharply arched. Alae well separated, thin
and pointed, often sharply. Sometimes the central or lateral alae are bifurcated.

Sigmas of a single size class. 45-60-66 ym (Fig. 5b). Arch shallow, may be even or eccentric.
Shaft moderately thick. Points very sharp with one bent out of the plane of the shaft.

Forceps of a wide range of sizes or more probably of a single size class, but may appear as multi-
ple size classes. 9-SO—87 ym (Fig. 5a, e). Shape highly variable, from V-shaped with legs nearly par-
allel to legs almost toxa-like. The most common form is long, slender, with legs nearly parallel. Spines
small but obvious on inner edge, all pointing upwards. Outer edge with few, somewhat blunt spines
with the exception of those on the upper edge where the legs join. Here the spines are erect, obvious
and sharp.

ETYMOLOGY. — This species was named macrostylosa in recognition of the large size of its
styles.

REMARKS. — This species is distinguished by the large size of its styles and tylotes and the ex-
treme size range of its forceps. In some respects it resembles Forcepia (Forcepia) topsenti Lundbeck,
1905, which has large styles and tylotes and forceps of a similar range of shapes. However, in all other
respects it differs. In F. topsenti, isochelae are of one size class, sigmas are significantly larger and sur-
face features are quite different from those detailed for the new species.

Forcepia (Forcepia) hartmani sp. nov.
Figs. 7 and 8

MATERIAL. — Holotype: CASIZ 53463, U.S.A., California, Monterey County, Pescadero
Point, 17 Mile Drive, April 28, 1982. Three pieces. Depth, intertidal. Coll. W. Lee. Paratypes:
CASIZ 5346S, U.S.A., California, Monterey County, Point Lobos, March 1984. Coll. D. Chivers and
W. Lee; CASIZ 35911, U.S.A., California, Sonoma County, Bodega Bay, Bodega Marine Labs, Au-
gust 9, 1983. Two pieces. Depth 0.5 m, rocky intertidal; CASIZ 017311, U.S.A., California, Farallon
Islands, Southeast Farallon Island, April 4, 1977. Coll. B. Bowman and C. Chaffee. Numerous pieces.

236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

Depth intertidal, +0.4 m under overhang; CASIZ 35961, U.S.A., California, Sonoma County, Bodega
Bay Marine Laboratory, August 9, 1983, open coast west of aquaculture building, Depth, intertidal,
-0.5 m, rocky intertidal. Coll. S. Ward and A. Miller. Other Material: CASIZ 18, CASIZ 31, CASIZ
3662, CASIZ 4593, CASIZ 6923, CASIZ 20358, CASIZ 31282, CASIZ 35961, CASIZ 53411,
CASIZ 53461, CASIZ 53464, CASIZ 59662, CASIZ 78254, CASIZ 108925, YPM 1540 and YPM
1697A.

DISTRIBUTION. — Present known range: Point Lobos, Monterey Co. to Bodega Bay Headlands
and Farallon Islands, California. Type locality: Pescadero Point.

HABITAT. — Rocky intertidal, in deep pools or rocky overhangs from about + 0.4 m to encrust-
ing rocks, -0.4—0.5 m.

SHAPE. — Thin to thick encrusting with a more or less flat surface. Incrustations up to 1.6 cm
thick. Size of largest piece of holotype 4.8 cm long x 4.2 cm wide x 1.6 cm high. Incrustations may
cover a much larger area.

CoLor. — In life, honey yellow, yellow gold, yellow, gold tan, buffy citron, buff; light tan in
ethanol.

OSCULA. — Oscula numerous, 1.0 to 4.0 mm across, round or somewhat irregular with slightly
raised membranous lips. In the thickest specimens the oscula tend to be within the surface grooves
where their shape and the membranous lips may not be easily seen.

TEXTURE AND SURFACE CHARACTERISTICS. — Consistency slightly compressible, friable. Sur-
face nodular and ridged, ridges somewhat hispid and delineating shallow grooves. In larger speci-
mens the grooves are deeper.

ECTOSOME. — (Fig. 8a) Exceedingly dense, crustlike. Made up of tightly bound masses of
tylotes to subtylotes parallel to the surface. Generally 85 to 91 1m in thickness but may reach over
200 pm in some places due to additional, looser, accumulation of tylotes underneath.

CHOANOSOME. — (Fig. 8a) A reticulation of wide tracts of styles with an overlying, looser, less
structured reticulation of random styles and tylotes. The tracts range in size from small, 24 to 28 um to
large, 60 to 72 pm. Within the choanosome are large strands of tissue with massive numbers of
microscleres, most notably sigmas.

MEGASCLERES. — Styles to subtylostyles of a single size class. 169-202-221-281 um x
7-8—9-10 1m (Fig. 7e). These vary from straight to strongly curved, the curvature occurring on the
upper 1/2 to 1/3 of the spicule. Most are simple styles but some may have tiny spines on either head or
tip. Even when some spines occur, the spicules look more like a normal style than an acanthostyle. The
shaft is equally wide throughout most of its length. The head is evenly and well rounded but may ap-
pear slightly swollen. The tip end often has a small indentation which temporarily reduces the shaft
width just prior to a long, sharp, tornote-like tip. The appearance is as though the shaft was pinched in-
ward before the tip, leaving a slight indentation.

Tylotes to subtylotes ofa single size class. 137—166—185—205 tum x 4—5—6 um (Fig. 7d). Mostly
straight to very slightly bent. Shaft of even width or slightly wider centrally. The heads are distinctly
tylote or strongly subtylote. There is a tendency for the swollen heads to be elongated, with nearly par-
allel sides. This is especially noticeable on the smallest spicules. Spicules with ends often unequal in
size.

MICROSCLERES. — Arcuate isochelae of a single size class. 18—23-34—38 wm (Fig. 7b). Thick
shaft with moderate curvature. Alae tend to be short, well separated, with rounded tips which may be
slightly flared. Lateral alae fused to shaft 3/4 of their length.

Sigmas ofa single size class. 30-42-48-55 um (Fig. 7c). Sigmas with low arch and tending to be
elongate with a relatively thick shaft. One end has a wider curvature than the other end and with a
somewhat curved, sharp point. The opposite end is narrower and more compact, with a very sharp and
sharply bent spine, usually bent out of the plane of the shaft.

LEE: CALIFORNIA FORCEPIA (PORIFERA)

FIGURE 7. Scanning electron micrographs of the spicules of Forcepia (Forcepia) hartmani sp. nov. a. Forceps 10,000x, b.
Isochela 3,000x, c. Sigma 2,000x, d. Tylote to substylote 700x, note the difference in the two ends, e. Style 500x.

FIGURE 8. Forcepia (Forcepia) hartmani sp. nov. a. Cross-section 40x, b. Holotype (CASIZ 053463), dimensions 4.8 x
4.2 cm wide and 1.6 cm high.

238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

Forceps ofa single size class. 5-8—9—11 jm (Fig. 7a). These are generally horseshoe-shaped with
a narrow to moderate range in the angle of the two legs from the median between them. The legs are
heavily spined, the spines all curved upward and well spaced. There are three rows of spines seen on
each side; one facing inwards, one along the middle of the leg and one on the outer edge. The tips of the
legs are widened and set off by three large spines.

ETYMOLOGY. — This species is named after the sponge systematist and biologist, Doctor Wil-
lard Hartman. Hartman introduced me to the marvelous world of sponges and has been a colleague in
producing the sponge chapter in the latest edition of the Light’s Manual (Lee, Hartman, and Diaz, in
prep.). Doctor Hartman has done much for our understanding of the systematics and biology of the
Porifera. He has likewise become a valued friend.

REMARKS. — This species originally appeared in material reviewed in Hartman’s 1975 descrip-
tion of Lissodendoryx firma where he described the occasional occurrence of forceps in this species.
This is a very understandable error, likewise initially made by this author, since the spicule comple-
ment of these two species is so similar. In fact, it was only by separating out all supposed L. firma with
forceps and comparing this group with those that had no forceps, that it became obvious that the for-
ceps-bearing specimens were clearly different from L. firma. While the spicule complement (other
than forceps) is superficially similar between the two species, the details of their structure differ sig-
nificantly. Most important are the obvious but subtle differences in skeletal structure, especially in the
nature of the ectosome and the details of the choanosomal tracts.

The ectosome of Forcepia (Forcepia) hartmani is exceedingly thick and tightly bound with
tylotes to subtylotes parallel to the surface but with few, if any, spicules penetrating the surface. In
Lissodendoryx, this area is made up of palisades of subtylotes that may be perpendicular, parallel or at
an angle to the surface. Spicule penetration of the surface and the formation of brushes is common.

The choanosome of Forcepia (Forcepia) hartmani is made up of obvious, bold, thick tracts that
form a reticulation over which can be found a looser, more random reticulation with many random
spicules. In Lissodendoryx firma there are distinct to vague tracts just under the ectosome. These are
far less dominant than those in Forcepia. Also, in Forcepia the choanosome is made up of a distinct re-
ticulation of thin tracts. The deeper one looks, the more random the reticulation appears.

While these differences are consistent, they nevertheless are subtle. However, even more subtle
is the occurrence of forceps. These microscleres are tiny in Forcepia (Forcepia) hartmani and are ex-
ceedingly difficult to find unless one is well aware that they may be present. Thus, the similarities be-
tween the two species can lead to a hurried, and incorrect, identification.

Forcepia (Forcepia) hymena (de Laubenfels, 1930) n. comb.
Figs. 9, 10, and 11

Wilsa hymena de Laubenfels, 1930 (Fig. 10)

MATERIAL. — Holotype: U.S.N.M. 21515, California, Monterey Co. Monterey Bay, May 9,
1929, Depth 700 m, Coll. E. F. Ricketts; Paratype: B.M. 29.8.22.62, California, Monterey Co.,
Monterey Bay, May 9, 1929, Depth 700 m, Coll. E. F. Ricketts. Reference specimen: BIC-SIO
P-1366, BIC-SIO P-1367, CASIZ 146075, R-12. San Diego Trough, California. 32°34.5’N,
117°33'W, 1170-1216 m, 25’ otter trawl. Mud. October 29, 1970, 1900-2147. R/V Agassiz. Coll. F.
Rokop, S. Luke.

DISTRIBUTION. — San Diego Trough to Monterey Bay, California.

HABITAT. — Possibly mud, 700-1216 m.

SHAPE. — Globular, massive; BIC-SIO P-1366, 5.3 x 2.0 x 3.5 cm. high. BIC-SIO P-1367, 2.1 x
1.3 x 0.8 cm. high, 1.7 x 1.5 x 0.6 high, and 3.0 x 2.0 x 1.4 cm high.

LEE: CALIFORNIA FORCEPIA (PORIFERA)

FIGURE 9. Scanning electron micrographs of the spicules of Forcepia (Forcepia) hymena (de Laubenfels, 1930). a. Forcep
5,000x, b. Forceps upper end showing fewer and smaller spines, c. Large sigma 500x, d. Small sigma 500x, e. Isochela 3,000x,
f. Isochela 4,000x, g. Substylote 400x, h. Style 200x.

FiGurE 10. Light micrographs of the spicules of de Laubenfels’ Wilsa hymena, 1930 (Holotype USNM 21515). a. Forceps

600x, b. Large sigma 500x, c. Small sigma 500x, d. Isochela 600x, e. Substylote 400x, h. Style 200x. Note: The quality of these
images was largely influenced by the poor condition of de Laubenfels’ slides.

240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

FIGURE |1. Forcepia (Forcepia) hymena (de Laubenfels, 1930). Reference specimen CASIZ 146075, dimensions 5.3 x
2.0 cm wide x 3.5 cm high.

COLOR. — In life, not recorded for reference specimen; holotype recorded as “pale drab.” Dark
tan to brown in ethanol.

OSCULA. — Numerous, round to somewhat elongate or irregular, 1.5 to 4.0 mm across. Flush
with surface or somewhat recessed, no lip or rim.

TEXTURE AND SURFACE CHARACTERISTICS. — Soft, compressible, spongy. Surface with some
areas superficially smooth but mostly slightly roughened, verrucose. Surface with shallow grooves
delineated by smooth ridges and small rounded conules.

ECTOSOME. — The ectosome is delineated by a dermal membrane, 48 to 72 um thick which con-
tains microscleres in abundance, especially the macro-sigmas. Below is a compact bundle of
subtylotes reaching up to 96 um in thickness. The subtylotes are tightly bound together and parallel to
the surface.

CHOANOSOME. — The choanosome is dominated by a rugged reticulation of styles between
tracts of styles. The larger tracts range from about 45 to 95 um thick. Smaller tracts of more loosely
bound styles may also be found, these ranging from 19 to 24 um. The entire choanosome appears as a
very rugged, almost random reticulation with more or less parallel tracts running through it. The
choanosome is also packed with sand grains and detritus. The combination of a soft, compressible tex-
ture with the presence of sand grains and detritus made it impossible to obtain good cross-sections.
Accordingly, the structure detail had to be pieced together from observations of numerous slides of
varying quality.

MEGASCLERES. — Styles of a single size class. 533-601-697 x 13-14-16 um (Figs. 9h, 10f).
Smooth, width even throughout. Most somewhat curved near middle. Tip tornote-like with sharp
point.

Subtylotes of a single size class. 273-327-448 x 5.5—6.9-7.0 um (Figs. 9g, 10e). Straight,
smooth, slightly wider near center. Ends mostly subtylote, or tylote with head somewhat elongate.
Frequently ends unequal, one subtylote, the other tylote, and of different sizes.

MICROSCLERES. — Arcuate isochelae of a single size class. 24-38-48 ym (Figs. Ye, f; 10d).
Strong arch with alae long and well separated. Frontal ala long, narrow and sharply pointed, often di-
vided into two or three small or independent alae. Lateral alae longer, attached to shaft by 1/2 to 3/4 of

LEE: CALIFORNIA FORCEPIA (PORIFERA) 24]

their length and curved toward the shaft with a narrowed but rounded tip. Lateral alae tend to have
hints of additional divisions, but these are never complete but seen only at the outer edge. Frequently,
one end differs from the other relative to such divisions.

Sigmas of two size classes. Small: 60-77-99 um (Figs. 9d, 10c). Somewhat elongate, with rela-
tively deep and eccentric arch. Both ends with tips moderately bent inwards in the same plane as the
shaft. Large: 169-208-243 um (Figs. 9c, 10b). Elongate with moderate arch. Arch not eccentric but
with one end with a rounder curvature than the other and a sharp point which is only moderately bent.
The other end with a point more obviously bent. Both points in the same plane as the shaft.

Forceps of a single size class. | 1-19-26 um (Figs. 9a, b; 10a). Legs long, parallel to about 2/3 the
distance from the tips where they are very slightly angled outwards. Wide spines which point up-
wards, mostly on the inner side of the legs. Many fewer spines on the outer edge. The legs terminate in
saucer-like caps. Spines on upper edge where the legs join, fewer and smaller than elsewhere.

REMARKS. — In 1930, de Laubenfels described a new genus and species, Wilsa hymena, from a
single specimen found on the macerated skeleton of a hexactinellid sponge. The specimen had an in-
tact ectosomal membrane containing abundant macro-sigmas. The choanosome was so enmeshed
with the hexactinellid on which it was residing that its structure could not be determined, although
some styles were found. Adjacent to the specimen and presumably contaminating it with its spicules,
was a specimen of Lissodendoryx kyma. De Laubenfels described Wilsa as containing styles, palmate
isochelae, macro-sigmas, sigmas of a smaller size class, and forceps. The forceps were described
through light microscopy as appearing smooth with only the faintest traces of spination. In addition to
these spicules, de Laubenfels noted the presence of some tornotes and arcuate isochelae.

In reviewing material from the Scripps Museum the author discovered in a mixed lot, several
pieces of a sponge that were obviously in the genus Forcepia. On preliminary examination it was
found that in many respects this material matched de Laubenfels’ Wilsa hymena. While it was possi-
ble to get reasonable cross-sections and scanning electron micrographs from the Scripps specimen,
this was not the case for the holotype of Wilsa hymena. The only material available were two de
Laubenfels microscope slides, one of which had little material on it. Fifty measurements were taken of
each spicule type for both the Wilsa type (U.S.N.M. 21515) and the Scripps material. The spicules
from these are compared in Table |. Note that the de Laubenfels measurements are designated by an
asterisk. Spicule widths for the de Laubenfels type are not given.

The palmate isochelae noted by de Laubenfels may well have been a contaminant. However,
most all of the isochelae seen on the de Laubenfels slide appeared to be arcuate. The arcuate isochelae
were thought to be a contaminant from Lissodendoryx kyma which de Laubenfels noted was living ad-
jacent to his specimen. Interestingly, the isochelae of L. kvma and those measured from the Scripps
specimens totally overlap in size range. Close comparison of the morphologies of the spicules that are
shared between the de Laubenfels type and the Scripps specimens show them to be very close or iden-
tical.

De Laubenfels erected the genus Wilsa for this single species, noting that it was most similar to
Esperiopsis forcipula, Lundbeck, 1905. Esperiopsis forcipula was later transferred to Leptolabis
(Topsent, 1904). In recent work for the Systema Porifera, Van Soest (pers. commun.) designated this
species as Forcepia (Forcepia) forcipula (Lundbeck, 1905). There is indeed a close resemblance be-
tween the two species, but it is clear that they are not the same. Among some of the differences, the
macro-sigmas of Forcepia (Forcepia) hymena are much larger and it has only a single size class of
isochelae, not two.

Given our present understanding of this group of sponges and the fact that the genus Wi/sa was
erected for a single, incomplete and contaminated specimen, Wilsa is hereby synonymized with
Forcepia and de Laubenfels’ holotype should be referred to Forcepia (Forcepia) hymena (de
Laubenfels, 1930). Since de Laubenfels’ holotype is both incomplete and contaminated, the reference
specimen may act as a subsidiary source of information on this species.

242 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

DISCUSSION

Five species of Forcepia are now known to occur in California. Four of these are newly described
herein; the fifth represents the assignment of Wilsa hymena de Laubenfels, 1930 to Forcepia and a
redescription based on newly found material. The family Coelosphaeridae Hentschel, 1923, to which
these species belong, may be generally distinguished by having an ectosomal tangential crust of
smooth diactinal (usually tylote) spicules, a reduced choanosomal skeleton composed ofa reticulation
of smooth or acanthose styles, with or without tracts and having sigmas, arcuate isochelae, and no
toxas. The genus Forcepia is the only forceps-bearing genus in the family. In addition, the ectosomal
spicules are always tylote or subtylote, the choanosomal spicules may be styles and may have
ectosomal spicules (tylotes or subtylotes) involved as well. In encrusting forms, one may sometimes
find a hymedesmoid structure replacing the reticulation, or elements of both. Van Soest (pers.
commun.) has used choanosomal skeletal structure to erect two subgenera. Species with a
hymedesmoid arrangement and acanthostyles with their heads embedded in a basal spongin layer are
placed in the subgenus Leptolabis. Those with styles or acanthostyles which are involved in a
choanaosomal reticulation are placed in the subgenus Forcepia.

In the genus Asbestopluma (family Cladorhizidae) some species also contain forceps but these
are structurally different than, and considered non-homologous with, the forceps of Forcepia. In addi-
tion, Asbestopluma differs from Forcepia in other significant ways. Asbestopluma tends to be abyssal,
with erect stalked growth forms and basal root adaptations. The upper part is penniform or with side
branches; the skeleton with a spicule axis divided into parallel fibers. Megascleres are styles or
subtylostyles in the axial and extra-axial skeleton and minutely spined tylostyles to tylostrongyles in
the coat of the stalk. Microscleres may be large, asymmetric palmate isochelae, sigmas, and forceps.

The California species of Forcepia are quite distinctive and can be readily separated.

KEY TO THE SPECIES OF FORCEPIA FROM CALIFORNIA

lias Monactsiare-acanthostylesa2-0.- chee iets cide ees Pec its | esome tee Forcepia (Forcepia) acanthostylosa sp. nov.
lib} Monactsare/styles;subtylostyles'or tylostyles: =| = «548-534 4.505. 9) 06 5 9 cele) eee 2
2a: Isochelaciof2)distinctisizeiclasses:..: «Givi « eid ;oed ie Soeie © cp cue eeey rs, od ee cen key ters an meee 3

2b. Isochelae of 1 distinct size class

3a. Tylotes small to moderate, 222-335 pm. Forceps of 2 distinct size classes, 11-28 um, 36-SSpm............
Sc ye pone eas ON Toms hcl OMT PN aT Re tel enO mene tal co iotis aaah Forcepia (Forcepia) elvini sp. nov.

3b. Tylotes large, 309-540 ym. Forceps either of many size classes or of wide range,9-87 um ...............
ee ee arc ia arnt oie ey ta aa Me te Shee WES MY Ree RE Forcepia (Forcepia) macrotylota sp. nov.

4a. Styles small, 169-281 pm. Sigmas | sizeclass.................. Forcepia (Forcepia) hartmani sp. nov.
4b. Styles large 533-697 um. Sigmas of 2 size classes, the larger being of exceptional size 169-243 pm ..........
ee a, tn Ber ee oe a oe ke oe Forcepia (Forcepia) hymena (de Laubenfels, 1930)

Of the species discussed, most need no further explanation as they clearly possess the usual char-
acteristics of Forcepia and have no circumstances surrounding them that would complicate their taxo-
nomic placement. However two species, Forcepia (Forcepia) hartmani and Forcepia (Forcepia)
hymena do merit further discussion.

As noted earlier, F. (Forcepia) hartmani was originally described as a forceps-bearing variant of
Lissodendoryx firma. Once forceps-bearing specimens were separated from those without, it was
clear that the two could be readily separated in other ways as well. Both genera are in the family
Coelosphaeridae and are closely related. In addition, their spicule complements other than forceps are
amazingly similar. In a like manner, while their skeletal structures are distinct, they are close enough
in detail to be confused if not examined carefully. To complicate these problems, the forceps in F.
(Forcepia) hartmani are.extremely small and not easily seen unless specifically sought for, or sub-

LEE: CALIFORNIA FORCEPIA (PORIFERA)

243

TABLE 1. Comparison of the spicule complement of Wilsa hymena de Laubenfels, 1930 (holotype) with
Forcepia (Forcepia) hymena (BIC-SIO 1366). All measurements in micrometers (um); measurements by

de Laubenfels (1930) with asterisk (*). Underlined = mean.

Morphological feature

Wilsa hymena
(holotype)

Forcepia (Forcepia) hymenia
(BIC-SIO 1366)

Smooth styles

(330-600 x 10—15)*

436-619-770 533-601-697 x 13-14-16
Subtylotes (not noted or 255-300-333 273-327-448 x 5.5—6.9-7.0
interpreted as tornotes)
Palmate isochelae 17—20* not seen
Arcuate isochelae in ectosome 19-26-39 24-38-48
Macro-sigmas approx. 250*
205-220-239 169-208-243
Sigmas J5-) 5)
55-75—108 60-77-99
Forceps 10—12*
faint spination with shallow spines
8-15-21 11-19-25
Acanthostyles (not mentioned) 47-53-62 not seen
Small styles (the larger with present not seen
microspined heads)
Dermal membrane 30—70* 48-72

jected to SEM analysis. Given these similarities, the geographic and depth distributions given for this
new species are probably incomplete. Accordingly, it will be important that current museum speci-
mens and newly collected material be carefully reviewed and the distribution data amended.

Forcepia (Forcepia) hymena is a prime example of the problems that can be generated by for-
mally describing a new species on the basis of a tiny fragment, which is admittedly contaminated. The
type of Wilsa hymena is sufficiently minuscule that further examination cannot take place without its
destruction, leaving it virtually useless. Only two slides were available of this material, with but one
being of any substantial use. It is always difficult to try to redescribe a species as poorly represented as
this one. However, extensive comparisons appear to strongly match those of the original material.
Noteworthy is the fact that Wilsa hymena as specifically described by de Laubenfels has never been
recorded since.

ACKNOWLEDGMENTS

The author gratefully thanks the David and Lucile Packard Foundation for their generous support
of the California Sponges Project. Without their insight and support none of this work could have oc-
curred. My deepest gratitude to David Elvin, Henry Reiswig and Paul Schroeder who all contributed
immensely with information, advice, critical reviews, and encouragement. Enormous thanks go to the
staff of the Departments of Invertebrate Zoology, and Entomology, at the California Academy

244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 18

of Sciences without whose help this work could have never been accomplished. Among those most
helpful were Bob Van Syoc, Chris Mah, and Elizabeth Kools in Invertebrate Zoology, and Darrell
Ubick in Entomology, who spent many long hours helping me with the SEM work. In addition to these
individuals I want to thank the Director of the Academy, Patrick Kociolek who generously facilitated
my use of the Academy SEM. I would like to also thank William Newman, Scripps Institution of
Oceanography and Director of the Invertebrate Museum for his assistance and for allowing free and
copious access to the Institution’s valuable collections. Finally, thanks go to others, including Jerry
Bakus, Eric Hochberg, William Austin, Willard Hartman, Klaus Rutzler, Rob Van Soest, and John
Hooper, all of whom contributed in so many important ways.

LITERATURE CITED

AUSTIN, W. C. AND B. OTT. 1987. Phylum Porifera. Pp. 6-29 in Marine invertebrates of the Pacific Northwest,
R. N. Kozloff, and L. H. Price, eds. University of Washington Press, Seattle.

CARTER, H. J. 1874. Descriptions and figures of deep sea sponges and their spicules from the Atlantic Ocean,
dredged up on board H.M.S. “Porcupine,” chiefly in 1869; with figures and descriptions of some remarkable
spicules from the Agulhos Shoal and Colon, Panama. Annals and Magazine of Natural History, ser. 4,
14:207-221, 245-257.

HADJU, E. AND J. VACELET. In press. Family Cladorhizidae de Laubenfels, 1936. /n Systema Porifera. A Guide to
the Supraspecific Classification of Sponges and Spongiomorphs (Porifera), J. N. A. Hooper and R. W. M.
Van Soest, eds. Plenum, New York.

HARTMAN, W. D. 1975. Phylum Porifera. Pp. 32—54 in Light’s Manual: Intertidal invertebrates of the Central
California Coast, 3rd ed., R. I. Smith and J. T. Carlton, eds. University of California Press, Berkeley.
HENTSCHEL, E. 1923. Erste Unterabteilung der Metazoa. Parazoa. Einziger Stamm und einzige Klasse der ersten

Unterabteilung: Porifera = Schwaémme. Handbuch der Zoologie 1:307-417.

Hooper, J. N. AAND R. W. M. VAN SOEST. In press. Systema Porifera. A Guide to the Supraspecific Classifica-
tion of Sponges and Spongiomorphs (Porifera). Plenum, New York.

LAUBENFELS, M. W. DE. 1930. The Sponges of California. Stanford University Bulletin 5(98):24-29.

. 1932. The marine and fresh water sponges of California. Proceedings of the United States National Mu-
seum 81:1—140.

LEE, W., D. ELVIN, AND H. REISWiG. In manuscript. The Sponges of California. n.p.

LEE, W., W. HARTMAN, AND C. DIAZ. In press. Phylum Porifera. /n Light’s Manual: Intertidal Invertebrates of the
Central California Coast, 4th ed, J. T. Carlton, ed. University of California Press, Berkeley.

LEVINSEN, G. M. R.1886. Kara-Haverts Svampe (Porifera). Dijmpha-Togtets zool. bot. Udbytte 341-372, pl.
XX1X—XXX.

LUNDBECK, W. 1905. 2. Porifera (Part II1). Desmacidonidae (Pars.). The Danish Ingolf Expedition, vol. 6,
pp.!—219 (Bianco) Luno, Copenhagen.

SOEST, R. W. M. VAN. In press. Family Coelosphaeridae Hentschel, 1923. /n Systema Porifera. A Guide to the
Supraspecific Classification of Sponges and Spongiomorphs (Porifera), J. N. A. Hooper and R. W. M. Van
Soest, eds. Plenum, New York.

TOPSENT, E. 1904. Spongiaires des Acores. Résultats des Campagnes Scientifiques accomplies sur son Yacht par
Albert ler Prince Scuverain de Monaco 25:1—280.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
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PROCEEDINGS
OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 19, pp. 245—395, 81 figs. October 26, 2001

The Green Lacewings (Neuroptera: Chrysopidae)
of Brazilian Agro-ecosystems

By
Sergio de Freitas
Universidade Estadual Paulista, Jaboticabal, Sao Paulo, Brazil
and

Norman D. Penny
Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

CALIFORNIA
ACADEMY
SCIENCES]

GOLDEN GATE PARK

SAN FRANCISCO
PUBLISHED BY THE CALIFORNIA ACADEMY OF SCIENCES

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 19, pp. 245-395, 81 figs. October 26, 2001

The Green Lacewings (Neuroptera: Chrysopidae)
of Brazilian Agro-ecosystems

by
Sergio de Freitas
Universidade Estadual Paulista, Jaboticabal, Sado Paulo, Brazil

and

Norman D. Penny
Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

Eighty-one species of Chrysopidae in six genera and two subgenera are described from
Brazilian agricultural systems. Forty-one of these are new species. They are: Nacarina
aculeata de Freitas and Penny (type locality: Birigui, S40 Paulo, Brazil); N. gladius de
Freitas and Penny (type locality: Birigui, Sao Paulo, Brazil); V. Javrasana de Freitas and
Penny (type locality: Jaboticabal, Sao Paulo, Brazil); N. sagitta de Freitas and Penny
(type locality: Ibitinga, Sao Paulo, Brazil); Ceraeochrysa dislepis de Freitas and Penny
(type locality: Itiquira, Mato Grosso, Brazil); C. dolichosvela de Freitas and Penny (type
locality: Jaboticabal, Sao Paulo, Brazil); C. squama de Freitas and Penny (type locality:
Jaboticabal, Sao Paulo, Brazil); Chrysoperla raimundoi de Freitas and Penny (type local-
ity: Jaboticabal, Sao Paulo, Brazil); Chrysopodes (Chrysopodes) adynatos de Freitas and
Penny (type locality: Itiquira, Mato Grosso, Brazil); C. (C.) copia de Freitas and Penny
(type locality: Jaboticabal, Sao Paulo, Brazil);C. (C.) crocinus de Freitas and Penny (type
locality: Birigui, Sao Paulo, Brazil); C. (C.) delicata de Freitas and Penny (type locality:
Itiquira, Mato Grosso, Brazil); C. (C.) elongata de Freitas and Penny (type locality: Luis
Antonio, S40 Paulo, Brazil); C. (C.) nigropicta de Freitas and Penny (type locality:
Itiquira, Mato Grosso, Brazil); C. (Neosuarius) karinae de Freitas and Penny (type local-
ity: Jaboticabal, Sao Paulo, Brazil); Plesiochrysa alytos de Freitas and Penny (type local-
ity: Itiquira, Mato Grosso, Brazil); Leucochrysa (Leucochrysa) bruneola de Freitas and
Penny (type locality: Itiquira, Mato Grosso, Brazil); L. (L.) catarinae de Freitas and
Penny (type locality: Fraiburgo, Santa Catarina, Brazil); L. (Nodita) affinis de Freitas
and Penny (type locality: Jaboticabal, S40 Paulo, Brazil); L. (N.) barrei de Freitas and
Penny (type locality: Itiquira, Mato Grosso, Brazil); L. (N.) confusa de Freitas and Penny
(type locality: Fraiburgo, Santa Catarina, Brazil); L. (N.) cornuta de Freitas and Penny
(type locality: Guaira, S40 Paulo, Brazil); L. (N.) forciformis de Freitas and Penny (type
locality: Itiquira, Mato Grosso, Brazil); L. (N.) furcata de Freitas and Penny (type local-
ity: Itiquira, Mato Grosso, Brazil); L. (N.) guataparensis de Freitas and Penny (type local-
ity: Luis Antonio, Sao Paulo, Brazil); L. (N.) ictericus de Freitas and Penny (type locality:
Birigui, S40 Paulo, Brazil); L. (N.) incognita de Freitas and Penny (type locality: Itiquira,
Mato Grosso, Brazil); L. (N.) interata de Freitas and Penny (type locality: Jaboticabal,
S4o Paulo, Brazil); L. (N.) lineata de Freitas and Penny (type locality: Luis Antonio, Sao
Paulo, Brazil); L. (N.) maculata de Freitas and Penny (type locality: Taquaritinga, Sao
Paulo, Brazil); L. (N.) michelini de Freitas and Penny (Jaboticabal, Sao Paulo, Brazil); L.
(N.) parallela de Freitas and Penny (type locality: Jaboticabal, Sao Paulo, Brazil); L. (N.)
retusa de Freitas and Penny (type locality: Balsamo, Sao Paulo, Brazil); L. (N.) robusta de
Freitas and Penny (type locality: Itiquira, Sao Paulo, Brazil); L. (N.) santini de Freitas

245

246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

(type locality: Jaboticabal, S40 Paulo, Brazil); L. (N.) scomparini de Freitas and Penny
(type locality: Itiquira, S40 Paulo, Brazil); L. (N.) squamisetosa de Freitas and Penny (type
locality: Birigui, Sao Paulo, Brazil); L. (N.) tabacinus de Freitas and Penny (type locality:
Itiquira, Sao Paulo, Brazil); L. (N.) tenuis de Freitas and Penny (type locality: Luis Anto-
nio, S40 Paulo, Brazil); L. (N.) vignisi de Freitas and Penny (type locality: Itiquira, Mato
Grosso, Brazil); and L. (N.) vittata de Freitas and Penny (type locality: Ribeirao Preto, Sao
Paulo, Brazil). Information about crops on which these species were collected is given when

known.
TABLE OF CONTENTS

IAtPOMUCHON 64.5 6c 6 cca oes oa a wv ewes o's a selene kale tle Seu Oe 248
Materials‘and) Methods... «ca. . vai. assualt en ae ee eee eee ee 250
Crop Associations .....-..2+5.5.>5 ostnge Cee = oe 251
Systematic Treatment sys sced Sc ve aeea ls cisteeh ee a a emits aera a ee 252
Belonopterygini ... 23.5.8 k Sete RE Rae o ec ce 259
INQCOT WAG oa. 55 55.205 Ge eh nt Shad da ROMER a SUE SET Pe 254
Chrysopinit 3 202.228. SUeai es Po PON SNRs Oe Ee eee 258
@eraéochrysa OO MP! PE et ce orn 0 oe 259
Chrysoperla: ie one ns oc ok aa he oe Ee ee eee et ng on oe 269
ChivsopOdes (CHIVSODOGCS)) cca nek ec gas eyes tgs ores, hg a > 271
Chrysopodes (N€OSUGTIUS)) 5 oo.0. 5.2.6. 0.600%, ofa tgnyle «psn wh ech She panies) eh <r 278
PIESIOCHRY Sg 505, sisca%e uapnduendegedinGehs 2% <fiTy oof insaeleieeinte alee eee 279
LeuCOChtySititys ».jcroroc% Hers 24a oh aieeieons'3- eee ote? 2aa bee 281
Leucochrysa (Leucochrysa@) \.as nansies Saseenn 8) LS). ee Oe eee 282
Leucochrysa(Nodita)s 2.22 9728 O) SORA LD A, Oe ee 296
Tlustratignis!s 086 Soi6a7't 22 yaa Oh) DM OPA OTE 1 BS DU 313
Acknowledanicnts: Pees ee eee a een ae eee 4. oat 395
Miteratne Cited ey ee cure aie tne eee ete Se tus oes Gye Eaten Gia. Bi)

Brazilian agricultural production is among the largest and most diverse in the world, generating
about $777 billion annually in foodstuffs and fiber and accounting for 8% of total gross domestic
product (Ministério de Agricultura, 1999). Agriculture is also the largest source of export revenues for
the country, producing $11.788 billion annually, or 24.55% of all export products (idem). For exam-
ple, Brazil is the largest producer of coffee, orange juice, and sugar in the world, and is second only to
the United States in soybean production. It has been said that the weather in Sao Paulo will materially
affect futures markets in Chicago and New York for such commodities as soybeans, coffee, and or-
ange juice. A more colorful way of phrasing this interdependence is that when a farmer sneezes in Sao
Paulo, a trader in Chicago catches a cold.

To protect these crops against pest damage a variety of strategies have evolved. As in North
America, there is a growing desire to rely less on pesticides and increase use of natural predators and
parasites. Two components of natural predator protection are inundative releases and oviposition aug-
mentation. However, in order for either of these strategies to be successful it is necessary for research-
ers to know what species of predators and parasites can exist in these ecosystems and how they interact
with each other. A first step in this direction is to document the species of predators and parasites
which naturally coexist with pests in the fields, orchards, and plantations.

in the United States, the most frequently utilized natural predators for inundative release pro-
grams are members of the green lacewing genus Chrysoperla (M. J. Tauber et al. 2000), which as

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE)

PLaTE A. Adult Ceraeochrysa cincta on sorghum. B. Adult Ceraeochrysa caligata on rubber tree leaf. C. Adult
Ceraeochrysa caligata on rubber tree leaf. D. Third instar of Chrysoperla externa feeding on aphids.

248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

adults feed on grass pollen and naturally occur most frequently in grasslands. Other green lacewings
are more frequently found in orchards (Ceraeochrysa spp.) and forests (Leucochrysa spp.).

Green lacewings often are voracious and aggressive predators as larvae, being quite effective in
feeding on phytophagous insects under ideal conditions. Neumark (1952) found that female
Chrysoperla carnea (Stephens) in Israel will lay up to 679 eggs. Smith (1921, 1922) counted up to 617
eggs per female of Chrysopa oculata Say in North America. Withycombe (1923) recorded larval
Chrysoperla and Chrysopa in Europe feed on an average of about 100 aphids during their develop-
ment, while Matsuda (1928) recorded as many as 900 aphids or 3,780 scale insects for each larva of
Japanese species. Neumark (1952) fed Chrysoperla carnea 6,487 scale insect eggs during the 14 day
cycle of its larval development. Larval green lacewings are known to feed on adult aphids
(Aphidoidea), scale insects (Coccoidea), mealy-bugs (Pseudococcidae), saw-fly larvae (Diprion-
idae), mites (Acari), Lepidoptera larvae and eggs, syrphid fly larvae (Syrphidae), dusky-wing larvae
(Coniopterygidae), thrips (Thysanoptera), and small Coleoptera larvae. However, although known as
general predators, there often is a strong link between host plant and associated species of green lace-
wing indicating a more specific association with prey items than is generally believed.

Brazil produces a tremendous number of crops in many different vegetation structures from open
fields of gramineous crops (barley, maize, oats, wheat) to vines (grapes, guarana), to tree crops (or-
anges, coffee, cacao), to selective harvesting of forest products (rubber, cupuacu). These crops often
contain a rich assemblage of chrysopid species, some of which have never been formally described.
Previous studies have almost always listed chrysopids as Chrysopa sp., a genus which is now re-
stricted to the Northern Hemisphere. Because of the rapidly evolving state of chrysopid taxonomy, all
previous identifications must be treated with a great deal of caution. Formal names and descriptions
are herein provided to facilitate studies of species biologies, interactions, and the role each species
plays in protecting these food plants. This monograph 1s primarily intended for use by taxonomists at-
tempting to identify species collected in these agricultural systems. Other crop protection specialists
may be able to identify the included species, but this should be done with caution. The male and fe-
male genitalic sclerites and membranes illustrated herein are tiny (often much less than one mm in
length) and require careful dissection and staining to observe clearly. A considerable amount of expe-
rience is needed to confidently recognize these structures properly.

MATERIALS AND METHODS

For this study 579 adults were collected by the senior author and his students on nine crops from
35 different localities in seven different states using several different methods throughout the year.
These methods included sweep net sampling, lights, flight intercept traps (Malaise traps), and Molas-
ses traps. Molasses traps were small plastic water bottles suspended from trees. Each bottle had a 10%
molasses solution inside and two 2 cm holes cut in the sides of the bottle. Chrysopids would enter
through the holes and could not find their way back out. While the number of crops and number of lo-
calities collected during this study was not comprehensive, in a country the size of Brazil the number
of potential crops and collecting localities is enormous. It was felt that enough information had accu-
mulated that this monograph was warranted. Certainly, future collecting on additional crops and in ad-
ditional localities will reveal further information about these and perhaps other species.

Many specimens were pinned and others were preserved in 75% ethyl alcohol. To study the
wings, they were detached from the body and mounted dry on microscope slides. Abdomens were de-
tached, macerated in warm 10% KOH, then stained with Chlorazol Black E. After study, wings were
glued to a small card on the pin below the specimen and abdomens were placed in small vials of glyc-
erin below each specimen. Drawings were made using dissecting and compound microscopes with
camera lucida attachments. Inked drawings were then scanned, using Abode Photoshop, and arranged
into composite plates.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 249

This study relies much more heavily on elements of the female genitalia than most previous stud-
ies. We have found that the form of the spermatheca (vela and ventral impression), spermathecal
ducts, subgenitale and surrounding area are much more diverse and reliable for species recognition
than previously thought.

Types of designated new species will be deposited in the Museu de Zoologia/USP, Sao Paulo, SP,
Brazil. Other material examined specifically for this study is deposited in the Sergio de Freitas Collec-
tion, Departamento de Fitossanidade, Universidade Estadual Paulista, Jaboticabal, Sao Paulo (SDF),
unless specifically indicated as deposited at California Academy of Sciences (CAS), San Francisco,
California, U.S.A.

CROP ASSOCIATIONS

Specimens were collected and examined from the following crops: rubber, orange, corn (maize),
guava, apple, cotton, cashew nut, cantaloupe and eucalyptus.

The insects collected in rubber tree (Hevea brasiliensis) plantations were identified as members
of 42 species in 4 genera, as follows: Leucochrysa (Leucochrysa) ampla, L. (L.) pretiosa, L. (L.) varia,
L. (L.) bruneolus; L. (Nodita) affinis, L. (N.) camposi, L. (N.) cruentata, L. (N.) heriocles, L. (N.)
lancala, L. (N.) lateralis, L. (N.) marginalis, L. (N.) marquezi, L. (N.) melanocera, L. (N.) rodriguezi,
L. (N.) barrei, L. (N.) forciformis, L. (N.) furcata, L. (N.) guataparensis, L. (N.) ictericus, L. (N.) in-
cognita, L. (N.) interata, L. (N.) retusa, L. (N.) robusta, L. (N.) scomparini, L. (N.) tabacinus, L. (N.)
vignisi; Ceraeochrysa cincta, C. caligata, C. claveri, C. cubana, C. everes, C. sanchezi, C.
tenuicornis, C. pennyi; Plesiochrysa elongata, P. alytos; Chrysopodes adynatus, C. delicata, C.
nigropicta, C. polygonica; Chrysoperla externa, and C. defreitasi.

In orange groves (Citrus sinensis) the following species were encountered: Leucochrysa (N.)
camposi, L. (N.) clepsydra, L. (N.) cruentata, L. (N.) walkerina, L. (N.) lancala, L. (N.) rodriguezi, L.
(N.) affinis, L. (N.) ictericus, L. (N.) maculata, L. (N.) michelini, L. (N.) vittata; Chrysopodes
lineafrons, C. polygonica, C. spinella, C. divisa; Nacarina pletorica, N. lavrasana; Plesiochrysa
brasiliensis; Ceraeochrysa cincta, C. caligata, C. claveri, C. cubana, C. dolichosvela, C. everes, C.
paraguaria, C. sanchezi, C. scapularis, C. tucumana, C. citrinus, C. pennyi, Chrysoperla externa, and
C. raimundoi.

In corn fields (Zea mays) the following were found: L. (N.) lancala, L. (N.) cornuta, L. (N.)
ictericus, L. (N.) rodriguezi, L. (N.) squamisetosa; Chrysopodes indentata, C. crocinus, Nacarina
panchlora, N. aculeata, N. sagitta, N. gladius; Plesiochrysa brasiliensis, P. elongata, P. alytos;
Ceraeochrysa cincta, C. claveri, C. cubana, C. everes; and Chrysoperla externa.

Guava (Psidium guajava) yielded the following species: L. (N.) camposi, L. (N.) cruentata;
Chrysopodes lineafrons, C. karinae; Plesiochrysa brasiliensis, P. alytos; Ceraeochrysa cincta, C.
claveri, C. cubana, C. everes, C. montouana, C. paraguaria, C. sanchezi, C. squamma; Chrysoperla
externa, and C. raimundoi.

The following species were found in apple (Malus pumila) orchards: Leucochrysa (L.) boxi, L.
(L.) catarinae, L. (N.) intermedia, L. (N.) confusa; and Chrysoperla externa.

Cotton (Gossypium spp.) fields yielded the following species: Leucochrysa (N.) interata, L. (N.)
parallela,; Ceraeochrysa cincta, C. cubana; and Chrysoperla externa.

In cashew nut (Anacardium occidentale) plantations the following species were found:
Leucochrysa (N.) rodriguezi and Chrysoperla externa.

The following species were found in cantaloupe (Cucumis melo) fields: Ceraeochrysa sanchezi
and Chrysoperla externa.

Eucalyptus plantations contained the following species: Leucocrysa (N.) aurantiacus, L. (N.)
guataparensis, L. (N.) lineata, L. (N.) robusta, L. (N.) santini, L. (N.) tenuis; Chrysopodes divisa, C.
elongata; Ceraeochrysa cubana, C. pennyi, C. scapularis; and Chrysoperla externa.

250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

SYSTEMATIC TREATMENT

The identification of Neotropical Chrysopidae is difficult. Of the 489 species described from the
New World south of the U.S.A. border, 290 of them were described by Padre Longinos Navas during
the first third of the 20th century (Penny 1978). His descriptions were brief, often not diagnostic, and
usually not accompanied by useful illustrations. To complicate matters, he sometimes placed type
material in his private collection, much of which has subsequently been destroyed. To understand the
species he described, it is necessary to visit many of the European museums where types exist, as well
as study the remnants of the Navas collection. The late Phillip A. Adams studied almost all of Navas’
remaining types, frequently macerated and stained the abdomens, took notes, and made pencil
sketches of male and female genitalia. These notes and sketches are now in the Entomology Depart-
ment of the California Academy of Sciences and have proven invaluable in the preparation of this
manuscript. In the individual species descriptions of this monograph, where no indication is given that
types have been studied, the species have been identified using Adams’ notes and sketches of the type
specimens. Additionally, in 1992 the junior author received an Ernst Mayr grant from the Museum of
Comparative Zoology to study Banks and Schneider types at that museum. A set of specimens from
the California Academy of Sciences was compared with types at Harvard University at that time and
has provided valuable comparative material. Less detailed studies have been made by the junior au-
thor of some Navas types in the British Museum (Natural History) (London) and Muséum national
d’Histoire Naturelle (Paris) but have not been included in the material examined section of species de-
scriptions.

Three publications have also exceptionally useful for interpretation of the Neotropical chrysopid
fauna. Adams and Penny (1987) covered not only part of the region under consideration, but also one
of the two main tribes involved in the present study. This paper provides keys to all species and in-
cludes known synonymy. Adams (1982a) described one of the major genera under discussion and
listed the 27 species known at that time, along with considerable synonymy. Finally, Brooks and
Barnard (1990) have provided a modern generic framework for many disparate species previously
placed together in the genus “Chrysopa.”

The most problematic group remains Leucochrysa (Nodita). This subgenus contains over 200 de-
scribed species, and no modern treatments of the individual species exist. Thus, we have relied almost
exclusively on the type specimens that we have seen and Phillip Adams’ notes and drawings for spe-
cies identification. The key included here is the first key of any Leucochrysa (Nodita) species in more
than 55 years. :

Weare following the classification and terminology used by Brooks and Barnard (1990) in their
world generic revision of Chrysopidae. We have included all species that we have found in agricul-
tural production areas, but have not intended this to be a complete treatment of all green lacewings
found in Brazil. Numerous other species are to be found in natural ecosystems and these probably oc-
casionally will stray into the more disturbed agricultural ecosystems. We hope eventually to publish
monographic revisions for the different tribes and genera of the Neotropics, but we intend this treat-
ment to be used for the specific habitat types indicated until more comprehensive studies can be com-
pleted. For the Amazonian region, a systematic treatment of Chrysopini is available (Adams and
Penny 1987).

Since we are relying so strongly on elements of the male and female terminalia for identification,
a few words of explanation of these structures are merited. A full complement of sclerites at the apex
of the male abdomen would include: a small quadrate sclerite directly below the anus (subanal plate),
followed in order by a thin arched structure (tignum) usually with a small medial projection (acumen);
an arched structure (gonarcus) with broadened apices (lateral arms), sometimes with a dorsal plate
(dorsal hood), a pair of dorsally projecting lobes (gonocornua), or caudo-ventrally projecting lobes
(entoprocesses) which partially wrap laterally around a membranous apical sac (gonosaccus); the

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 251

gonarcus usually has a caudally projecting medial lobe (mediuncus) which ends in medial point
(arcessus) which often is tripartite and decurved; below (and not connected to) the gonarcus and above
the gonosaccus there may be a long medial sclerite (pseudopenis); the gonosaccus often bears setae
(gonosetae) or small toothed plates (gonocristae) on its surface; lateral to the gonosaccus in
Belonopterygini there may be an elongate, pointed sclerite (paramere); finally, there may be a sclerite
above the fused eighth and ninth sternite (gonapsis), which in New World species takes the form of a
long rod. All elements of the male genitalia are never present, and the combination of presence or ab-
sence of these components is the best available criterion for generic recognition.

Female genital elements include a spermatheca consisting of heavily scleritized vela with a ven-
tral impression at its anterior end and connected near the mouth of this impression is a long duct
(spermathecal duct) ending in an appendix-like projection which is often hairy (see Fig. 30). At the
posterior end of the vela is a constricted and less sclerotized duct (bursal duct) which connects to an
expanded, membranous copulatory bursa. Two membranous accessory glands (bursal glands) empty
into the bursa. Above the bursa is a second set of membranous accessory glands (colleterial glands),
which terminate separately between the lobes of the lateral gonapophyses. On the body surface below
the lateral gonapophyses and above the seventh sternite is a sclerite (subgenitale) with paired dorsal
lobes and ventromedial lobe. In the Belonopterygini, the subgenitale can be protracted by an elongate,
membranous stalk. The ventromedial lobe may be quite small, or occasionally quite elongate. Often,
in the middle of this ventral lobe is a small, more heavily sclerotized indentation (crumena). In
Belonopterygini at the caudal apex of the seventh sternite there is a small medial sclerite
(praegenitale) with a central crumena. This probably reflects a migration and separation of the apex of
the subgenitale ventromedial lobe.

At present all known green lacewings found in agro-ecosystems in Brazil belong to three tribes,
all in the subfamily Chrysopinae. Adults can be distinguished using the following key.

KEY TO THE TRIBES OF CHRYSOPIDAE FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

1. Adults thick-bodied with prothorax wider than long (Fig. 1A); antennomeres wider than long beyond the first few

basal segments; larvae associated with antnests.. 2... 2... ee es Belonopterygini
1’. Adults thinner-bodied with prothorax usually as long as wide (Fig. 9A); antennomeres longer than wide beyond the first

few basal segments; larvae free-living on vegetation, often with bodies covered with debris and skins of prey . . . . 2
2. Forewings bearing dark spot at base of pterostigma; outer gradate series of crossveins at narrow angle (less than

45°) to pseudomedial vein (Fig. 41B); antennae longer than wings. .........--.....--.. Leucochrysini

2'. Forewing pterostigmal area completely pale; outer gradate series of crossveins at broad angle (greater than 60°) to
pseudomedial vein (Fig. 37B); antennae shorter than wings .........-...-.----2-2--2005- Chrysopini

TRIBE BELONOPTERYGINI NAVAS, 1913a

These green lacewings, when first seen, are remarkably robust. They are wider bodied than other
chrysopids. The pronotum is always wider than long and antennal segments are wider than long (Ad-
ams and Penny 1987). However, the species found in Brazil are among the least robust of the tribe and
some species can be confused with members of Chrysopini. Other species in the region, however, are
large and thick-bodied. We have seen some females from Rio Doce in Minas Gerais that are so big that
they can momentarily appear similar to some of the large, yellow megalopterans of the genus
Chloronia.

The tribe is worldwide in distribution. Most of the Old World species are in the genus /talochrysa
(Tjeder 1966). The dominant New World genus is Nacarina and all known Brazilian species are in
this genus. There are two additional New World genera (Adams 1978): Belonopteryx is a monotypic
genus with long, narrow wings known from only three specimens from Argentina; and Abachrysa is a

252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

genus with dark wings and dark maculations on the wings found in southeastern U.S.A. and the Carib-
bean (Brooks and Barnard 1990).

All known species of Belonopterygini are ant nest associates as larvae (Brooks and Barnard
1990). Their main contribution to agricultural ecosystems is probably to reduce the numbers of certain
species of ants in the fields. Their more frequent occurrence in corn fields may be an indication of
preference for ants living in a more open habitat.

Genus Nacarina Navas, 1915
Brooks and Barnard (1990) reported 14 species of Nacarina, all restricted to the New World with
11 of them found in South America. During this study, three of the 11 South American species were

seen, as well as four undescribed species. Adults can be distinguished using the following key.

KEY TO THE SPECIES OF NACARINA FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

J. Apical/segment of maxillary palps'swollen, bulbous (Fig..1D))........ 2. 2. «+ is) 2% Jo =e 2
1’. Apical segment of maxillary palps tubular, apically tapering (Fig. 3A)... . 2... 2... eee ee ee eee 5
2. Head and pronotum completely pale, without markings (Fig.2A).................+.-..... N. pletorica
2'. Vertex and gena with red marks; pronotum with lateral red stripe (Fig 1A)................. N. panchlora
3. Head without dark markings; lateral arms of gonarcus with funnel-shaped cavity (Fig. 7C) .......... N. sagitta
3'. Head with some dark markings (Fig. 5A); lateral arms of gonarcus without cavity (Fig. 4F)............... 4
4> Forewing inner and outer gradate veins paralleli((Fig-5B)5 =.= =|) 2) 262) ) ee 5
4Forewing inner and outer gradate veins divergent (Fig. 3B)) 25-22-5245: 3-55 os eee N. wagneri

5. No red markings on central raised area of vertex (Fig. 4A); parameres adherent to gonosaccus (Fig. 4G); gonosaccus

withinumerous:smalliponocristae\(Figs 4G), 5 2 ts es ee ee ed N. aculeata
5'. Two crescent-shaped red marks on raised central area of vertex (Fig. SA); parameres adjacent to gonosaccus (Fig. SF);
gonosaccus) withifew long sctae; no small) conocristae (Figs5E)- . 22 5 2. 3. 2 = 5 6

6. Intramedian cell quadrate (Fig. 6B); gena red; maxillary palps pale (Fig. 6C); inner margins of gonocornua form

Veshaped (HISSGE) ite cues ceca seatue sais dese ysiaa fal sy .ah erase ee asa eran ee N. lavrasana
6’. Intramedian cell triangular (Fig. 5B); gena pale; maxillary palps dark (Fig. SC); inner margins of gonocornua form
Weshaped: (Bigs) rsa ec a ca to ho Se 5 cid or cyan tiers TiS Jey opto 1A Seo eK oe N. gladius

Nacarina panchlora (Gerstaecker, 1888)

DIAGNOSIS. — This is one of two species with bulbous maxillary and labial palpi, short faces,
and broad bodies. There also appears to be the beginnings of a second intramedial cell. Nacarina
panchlora has much more red coloration on gena and pronotum than N. pletorica. The lateral
digitiform lobes of the female genitalia also appear distinctive. So far, we have only found this species
associated with corn fields.

HEAD. — Yellow and green. Vertex yellow, rugose, posteriorly elevated; two small, red
postocular spots at hind margin. Frons yellow with red marks below antennal bases. Clypeus and
labrum green. Gena red (Fig. 1C). Maxillary and labial palpi pale, last segment basally bulbous, api-
cally rounded (Fig. 1D). Antenna pale, first segment of flagellum larger than others.

THORAX. — Green with yellow mid-line. Pronotum twice as wide as long; anterior margin
arched; brown stripes laterally (Fig. 1B). Meso- and metanotum unmarked. Tarsal claws elongate
without basal expansion. Wings: Forewing green, with anterior endings of Radius (R)-Radial Sector
(Rs) and gradate veins dark. Subcosta (Sc) and Rs veins swollen. Intramedian cell (imc) quadrangular.
incomplete second intramedian cell. Partial third gradate series present (Fig. 1B). Forewing length
(2.55 cm): width (0.92 cm) =ratio (2.77). Hindwing length (2.28 cm): width (0.82 cm) =ratio (2.78).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 253

ABDOMEN. — Green with yellow mid-line. Female: Subgenitale with central projection (Fig.
1G). Spermatheca short, membranous posterior part folded upon itself. Bursa copulatrix acute anteri-
orly, with pair of accessory glands (Fig. 1E). Lateral to subgenitale a membranous lobe with two flat
pockets (Fig. 1 F).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Birigui, 6 April 1996, Scomparin, C. H. J.
(12)(corn); 3 September 1996, Scomparin, C. H. J. (12)(corn); Jaboticabal, 11 August 1991,
Scomparin, C. H. J. (1 2)(corn); Franca, 19 October 1994, Montovani, R.C. (1 )(corn).

Nacarina pletorica (Navas, 1919)

DIAGNOSIS. — This is one of two species with quite broad bodies, short faces, and bulbous apices
of palps. Nacarina pletorica can be separated from N. panchlora by the lack of genal and pronotal red
coloration and somewhat narrower wings, without partial third gradate series.

HEAD. — Pale yellow. Gena, clypeus, labrum, frons, vertex and antenna pale, without marks.
Face short (Fig. 2D). Maxillary and labial palpi pale, last segment large, bulbous, tapered to rounded
apex (Fig. 2C). Vertex rugose and raised posteriorly.

THORAX. — Pale yellow, unmarked. Wings: Venation pale, except anterior intersections of first
costal crossveins, basal subcostal crossvein, posterior endings of first R-Rs crossveins, gradates, ends
of Pseudomedius (Psm)-Pseudocubitus (Psc) crossveins, and apex of anal veins brown (Fig. 2B). Ime
quadrate. Pterostigma pale, unmarked. Rs, Psm and Psc swollen. Forewing slender; length
(17.8 mm): width (0.56 cm) = ratio (3.18). Hindwing green, except anterior ending of first costal
crossveins, posterior endings of first Rs-Psm crossveins, gradates, endings of Psm-Psc crossveins,
and endings of anal veins pale brown (Fig. 2B). Gradates, Psm, and Psc swollen. Hindwing slender,
length (1.6 cm), width (0.5 cm) = ratio (3.14).

ABDOMEN. — Yellow, unmarked. Ectoproct ventro-posterior angle tapering to sclerotized point.
Setae of callus cerci angled medially. Male genitalia: Eighth and ninth sternites fused; ninth smaller
than eighth and strongly tapered to apex; microtholi present (Fig. 2E). Gonarcus short and broad; lat-
eral arms short, with lateral lobe. Gonocornua plate-like with two apical lobes, the inner acute.
Arcessus short, apical lobe with medial hook flanked by short, rounded, lateral lobes. Gonosaccus
with many long setae (Fig. 2F, G).

MATERIAL EXAMINED. — BRAZIL: Sao Paulo: Jaboticabal, 18 December 1995, Franco, E.
(1c).

REMARKS. — The male specimen that we have available appears to have somewhat aberrant
wing venation in that the forewing has a second intramedial vein.

Nacarina wagneri Navas, 1924a

DIAGNOSIS. — Characterization was made from a faded, pinned specimen. This species is a
member of a species group having elongate, tubular, apical palp segments. The quadrate intramedian
cell separates N. wagneri from N. aculeata and N. gladius. The red head and pronotal markings sepa-
rate N. wagneri and N. lavrasana from N. sagitta, while the concave curvature of the inner gradates
separates N. wagneri from N. lavrasana.

HEAD. — Antennae pale. Red mark around antennal bases. Maxillary and labial palpi pale, last
palpimere slender, tapered and not inflated (Fig. 3A).

THORAX. — Pronotum twice as wide as long, with lateral red stripe. Mesonotum with lateral red
stripe. Wings: Forewing length (1.95 cm): width (0.69 cm) = ratio (2.83). Venation completely pale.
Intramedian cell quadrangular (Fig. 3B). Hindwing length (1.67 cm): width (0.60 cm) =ratio (2.78).

254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

ABDOMEN. — Pale. Female: Seventh sternite with apical knob (Fig. 3C). Spermatheca a large
chamber and several convoluted coils (Fig. 3E); accessory glands with huge sac which extends anteri-
orly to sixth segment (Fig. 3F). Subgenitale with medial projection with ventral cremena (Fig. 3D).

MATERIAL EXAMINED. — Brazil: Mato Grosso: Salabra, March 1940, collector unknown.

Nacarina aculeata de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museum de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled: “Birigui, SP, Fz. Sao Joaquim, 1 1/Jun/94, Scomparin, C. H. J., SP66” (on corn).

DIAGNOSIS. — This is a member of a species group having tubular, tapering palps. It has ovate
intramedian cell and dark palpi, which separates it from all other species, except N. gladius. Nacarina
aculeata can be separated from N. gladius by the shorter face, lack of red markings on the vertex,
shorter inner gradate series of the hindwing, shorter setae on ectoproct and sternite 8 + 9, shape and
longer length of the parameres, and spiny apex and dorsal margin of sternite 8 + 9.

The name “aculeata” comes from the Latin aculeatus meaning sharp-pointed. This name refers to
the field of spines along lateral margin of sternite 8 + 9 and the teeth at the apex of this sternite.

HEAD. — Golden yellow. Vertex, frons and clypeus glabrous. Face short. Gena pale, unmarked.
Vertex deeply depressed laterally near eye margin. Scape and pedicel with dorsal brownish red
stripes; basal segments of flagellum black striped on inner surface (Fig. 4A), more distal segments
fuscous, with setae black. Maxillary and labial palpi with black markings; last segment tapered and
slender. (Fig. 4C).

THORAX. — Yellow. Pronotum glabrous with lateral pale reddish brown stripe (Fig. 4A). Wing:
Venation pale, except R-Rs crossveins, Rs-Psm crossveins, Rs-inner gradate (Ig) crossveins,
gradates, Psm-Psc crossveins, marginal forked and unforked veins dark. Pterostigma unmarked.
Forewing length (1.6 cm): width (0.52 cm) = ratio (3.08). Intramedian cell triangular. Hindwing ve-
nation pale except costal crossveins, R-Rs transverse vein endings and gradates dark. Length
(1.38 cm): width (0.43 cm) = ratio (3.2) (Fig. 4B).

ABDOMEN. — Male: Densely covered by short and pale setae (Fig. 4D). Sternite 7 much larger
than sternite 8 + 9. Apex of ectoproct rounded (Fig. 4D). Sclerotized melanic band at anterior margin
of ectoproct with branch leading downward to melanic callus cerci, unlike the pale apodemes found in
all other species. Apex of sternite 8 + 9 with several heavily sclerotized teeth; subapical dorsal margin
of sternite with spiny lobe (Fig. 4E); without microtholi. Arcessus short, broad with dorso-medial pro-
jection terminating in decurved hook flanked by two rounded lobes (Fig. 4F, G, H). Lateral arms of
gonarcus with posterio-ventral portion extending below arcessus. Gonosaccus with few long setae;
ventral portion with field of numerous small gonocristae. Parameres parallel curved rods adherent to
gonosaccus (Fig. 4F).

MATERIAL EXAMINED. — Known only from the holotype.

Nacarina gladius de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled: “Birigui, SP, 6 April 1996, Scomparin, C. H. J.” (on corn).

DIAGNOSIS. — This is a member of a species group having tubular, apically tapered palps and a
triangular intramedian cell. It can be separated from N. aculeata by the presence of a double crescent
red mark on the vertex, much longer inner gradate series of the hindwing and characteristics of the
mate genitalia such as a more narrow arcessus, lack of spines on sternite 8 + 9 and lack of gonocristae
on the gonosaccus in N. gladius.

‘he name “gladius” comes from the Latin word for sword, and refers to the shape of the

parameres.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 255

HEAD. — Golden yellow. Vertex raised, rugose, with red double crescent marking. Scape and
pedicel with wide dorsal red stripe (Fig. 5A); first 37 flagellomeres with black stripe along medial
margin; more apical segments fuscous. Frons with red spot below antennal base and above anterior
tentorial pit. Gena black-banded near frons. Maxillary and labial palpi pale, apical segment tubular,
apically tapering (Fig. 5C).

THORAX. — Yellowish green. Pronotum with short and slender brown stripe at anterior angle;
posterior angle weakly red spotted. Meso- and metanota without markings (Fig.5A). Wings: Vena-
tion green, except costal crossveins, R-Rs crossveins and gradates dark (Fig. 5B). Pterostigma pale,
unmarked. Forewing length (2.23 cm): width (0.69 cm) = ratio (3.23). Hindwing length (1.8 cm):
width (0.6 cm) = ratio (3.0). Venation green, except middle costal crossveins dark.

ABDOMEN. — Green without marks. Ectoproct dorso-apical angle acute. Sternite 8 + 9 evenly
tapered, triangular; suture not visible, no microtholi (Fig. 5D). Gonarcus not vertically arched; lateral
arms ovate. Bases of gonocornua proximal to each other; inner margins U-shaped. Arcessus short,
narrow; with medial decurved hook and tiny rounded lateral lobes. Gonosaccus with few long
gonosetae. Parameres long, sword-shaped, conical, adherent to gonossacus.

MATERIAL EXAMINED. — Known only from the holotype.

Nacarina lavrasana de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP; Abril/93, Murtati, G.”

DIAGNOSIS. — This species can be separated from other South American species by the tubular,
apically tapered maxillary palpi and quadrangular intramedian cell. The only other species of
Nacarina in the region with these characteristics is N. sagitta, which has no red markings on head and
thorax. The male genitalia are also quite different, with the gonocornua of N. /avrasana much more
approximated basally and much shorter, and not having dorsal pockets of the lateral arms of the
gonarcus. This species also resembles N. cordillera from northern South America and Central Amer-
ica, but that species has the apex of male sternite 8 + 9 bluntly truncated.

The name of this species is derived from the town of Lavras in Minas Gerais State.

HEAD. — Yellow, marked with red. Vertex glabrous with red, double crescent-shaped marks
(Fig. 6A). Scape and pedicel with narrow red stripe dorsally; flagellum with medial dark stripe on
basal flagellomeres and completely brown apical ones. Gena with red spot bordering frons (Fig. 6C).
Maxillary and labial palpi pale, last segment slender and narrow, apically tapered (Fig. 6D).

THORAX. — Green. Pronotum with red lateral stripe. Meso- and metanota green without marks.
Wings: Venation green, except posterior endings of R-Rs crossveins and gradates of forewing dark.
Transverse vein between first and second median cells inflated (Fig. 6B). Forewing length (2.28 cm):
width (0.74 cm) = ratio (3.08). Hindwing length (2.02 cm): width (0.65 cm) = ratio (3.1).

ABDOMEN. — Green without markings. No microtholi. Male genitalia: Gonarcus arcuate, thick.
Bases of gonocornu proximal, apically decurved. Arcessus long, with small, rounded lateral lobes and
medial decurved hook. Cluster of setae on gonosaccus below gonocornu. Paramere slender, adherent
to gonosaccus (Fig. 6H, I). Female genitalia: Spermatheca short, extended as long wrinkled expan-
sion to bursa copulatrix. Spermathecal duct short, coiled. Two sets of colleterial glands long and fila-
mentous, extended to fifth segment (Fig. 6E, G). Subgenitale heavily-sclerotized, extended anteriorly
as sclerotized medial lobe (Fig. 6F).

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Sao Joao da Boa Vista, 12 October 1994,
Mello, S. R. C. (12 paratype)(orange); Jaboticabal, April 1993, Murtati, G. (1? paratype); Minas
Gerais: Lavras, 10 October 1990, Carvalho, C. F.

256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

Nacarina sagitta de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Ibitinga, SP; 19-12-88, Maia, A. M.”

DIAGNOSIS. — This species is a member of the group having slender, tapering pale palpi and
sharply tapered male sternite 8 + 9. It is close to Nacarina gladius, but can be differentiated by a suite
of characteristics: N. gladius has a triangular intramedian cell; gonarcus less vertically arched; bases
of gonocornua more closely approximated, lateral arms of gonarcus not inflated with internal pocket.

The name “sagitta” comes from the Latin word for arrow, and refers to the shape of the male
parameres.

HEAD. — Vertex, frons, clypeus, labrum and gena pale yellow. Antenna pale, unmarked.

THORAX. — Yellow. Pro-, meso- and metanota unmarked. Wings: Wing apex missing. Trans-
verse vein between first and second median cells and contiguous parts of Psc swollen. Intramedian
cell quadrangular (Fig. 7B).

ABDOMEN. — Apex of ectoproct truncate. Male genitalia: Dorsal apodeme of ectoproct forked
near base; ventral fork continues ventrally below ectoproct and terminates acutely (Fig. 7A). Sternite
8 + 9 partially fused with visible suture; microtholi present (Fig. 7A). Medial arch of gonarcus short
and strongly curved; lateral arms swollen with inner pocket which opens dorsally (Fig. 7C).
Gonocornua slender, with bases well separated. Arcessus elongate; apical lateral lobes as vertically
oriented plates and medial decurved hook. Group of little lobes below arcessus and laterad of
gonosaccus with long setae. Large, sclerotized, conical, apically arrowhead-shaped parameres extend
to arcessus (Fig. 7C, D).

MATERIAL EXAMINED. — Known only from the holotype.

TRIBE CHRYSOPINI SCHNEIDER, 1851

Chrysopini are the green lacewings most often seen in fields and on windows and store fronts at
night. Adults of some species, such as those in the Holarctic genus Chrysopa, can emit a foul-smelling
odor in self-defense. Adults of Chrysopini are small to medium-sized (8—20 mm forewing length) and
basically green in color with four similar transparent wings. Larvae of some genera, such as
Chrysoperla, have naked larvae, while larvae of other genera, such as Ceraeochrysa, cover their bod-
ies with debris and dried bodies of prey. As a group, they are indigenous to all temperate and tropical
parts of the world, except New Zealand.

Adult Chrysopini are morphologically conservative, so that the generic classification is based
primarily on sclerotized elements of the male genitalia. Adults of genera of Brazilian Chrysopini can
be distinguished using the following key:

KEY TO MALES OF CHRYSOPINI GENERA FOUND IN BRAZILIAN AGRO-ECOSYSTEMS
(modified from Adams and Penny 1987)

1. Tignum present (Fig::25E))<cicos0 S oc) See SOs SRS Se 2 ke S8 ae 2
[! Tignum absent:(Fig? 31 B) ic. A9.nnSecrct Secession? tenteas trerions task, domed ree geeree ts baie ae 3
2. Pseudopenis present; pronotum with four red spots or lateral stripes (Fig.37A,H) ............. Plesiochrysa
2’. Pseudopenis absent; pronotum without red marks (Fig. 24A,I)..............200 5-052 0 0s Chrysoperla
3. Gonapsis elongate (Fig. 8C); two hornlike structures on gonarcus or arcessus (Fig. 9E).......... Ceraeochrysa
3’. Gonapsis absent (Fig. 31B); hornlike structures absent from gonarcus-arcessus (Fig. 31E) ...........-.+.-. 4
‘. Mandibles blunt-tipped; wings with narrow costal area in most species (Fig. 30B)...... Chrysopodes (Neosuarius)

Mandibles with fanglike tip (Fig. 32E); wings with wide costal area in most species (Fig. 36B)
sh TAURI leat xi lallta te: Nate convo seas We eee eae oak ye cao ae ae er Chrysopodes (Chrysopodes)

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 251

Genus Ceraeochrysa Adams, 1982a

Adults of Ceraeochrysa are among the most colorful of all New World Chrysopini and most eas-
ily characterized. Individuals often have lateral or dorsal stripes on the scape and the pronotum usu-
ally has stripes or dark spots. Dark marks can also occur on the meso- and metanota. Dark margining
of veins can occur on the wing membrane and some species have swelling and darkening of veins at
the base of the posterior margin of the forewing. Females have a simple U-shaped spermatheca. Males
of all species have an elongate rod (gonapsis) above sternite 8 + 9. Adams and Penny (1987) recorded
the occurrence of a triangular gonapsis in one species of Chrysopodes. However, Brooks and Barnard
(1990) regarded a gonapsis as absent from Chrysopodes and we have never observed a gonapsis in
other species of Chrysopodes. Ceraeochrysa is the most speciose genus of Chrysopini in the New
World with more than 50 described species. We add an additional four species in the current treat-
ment. The genus 1s distributed from southern Canada (Penny et al. 1997) to northern Argentina (Penny
1978), although it is most abundant and speciose in the tropics. Ceraeochrysa cincta has been col-
lected on bananas and citrus in Honduras (Adams and Penny 1987) and must be considered a prime
candidate for biological control programs. In fact, many species of this genus are found in association
with orchards and probably contribute significantly to natural pest control. Lopez-Arroyo etal. (1999)
recommended three species of Ceraeochrysa as biological control agents showing great potential in
tropical America. Considerable recent work has been done on larval taxonomy of this genus by C. A.
Tauber et al. (2000) and Tauber and de Leon (2001). Fifteen species have been collected in Brazilian
agro-ecosystems and adults can be distinguished using the following key.

KEY TO MALES OF THE SPECIES OF CERAEOCHRYSA FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

[eeeeronotum with dark lateral stripes (Fig. LOA)... Ps.c caren ate) weno ye (drew sks aennen 2 2 nue apes Ohw selon 2
1’. Pronotum with two pair of dark circular marks (Fig. 15A); apex of sternite 8 + 9 with a pair of forcipate

Pam e eee FESO actin ir coir ae ke sec te) Sa. fone asc Ss mate ee Goan <h goin subg he Fed Gabe a C. paraguaria
2. Antennal scape without dark markings (Fig. 21A); spermatheca long (Fig.21C) ............ C. dolichosvela
Zee antennal scape. with stripes OL Spots (Bigs 1 VA) 502 5 2 3 52 wb se cus es ode © dyes so ye de evens 3
PE UNAl Reape wit lateral epots (hig. ISA): esecicspswa ¢ysyseeyt Qysxe Spe sfdeie ee ie apa isteem tas le Girels 4
Se eennanscane wvith dark stripes (Fig. 1 LA)... 2.5... 2ass y<, eres). 2)arrdye) an «loot, svmy ae Sinpin pawn, Socpsweun Bye Slanerace 5
4. Dorsal apodeme of ectoproct extended ventrally as apically-expanded projection (Fig. 8C).......... C. acmon
4'. Dorsal apodeme of ectoproct not extended ventrally as projection (Fig. 12C); gonarcus with dorso-medial,

POST SREP SCANT ea he) eee ee ee ee C. everes
Se oeane withidorse-medial dark stripe (Figs) 11A)\a ea) aft ser msi cast Sener Sie A remeeee 4 ew Sate Shel ew cree 6
Sede marnidorse-latcral dark stripe (Fig. LGA) 1. eccieds serteend a som, @ wl lewar lee dow anem day lank Gua vebershalh 9
6. Scape stripe not extended basally onto antennal fossa (Fig. 10A)..................2.2.00. C. cincta
6’. Scape stripe extended basally onto antennal fossa (Fig. LIA)... 22... ee ee 7
in ea emeraeclnnnodne (hiss lOA) =. 54. Se ST Lee ODP tm US Doe hae, C. caligata
Penn ureemee Minmaara(E isl A) rae ee i sine ee GP ho Lom er reras dl) gown Se 8
8. Scape with two dark stripes (Fig. 14A); spermatheca U-shaped (Fig. 14C); dorsal apodeme of ectoproct not

prolonged ventrally beyond ectoproct (Fig. 12C); gonapsis unforked(10F)................ C. montoyana
8’. Scape with single dark stripe (Fig. 12A); spermatheca not U-shaped (Fig. 10E); dorsal apodeme of ectoproct

extended ventrally beyond ectoproct (Fig. 11C); gonapsis forked caudally (Fig. 11F)............. C. claveri
POsnCnnal Hapeiuinipaled ero 2A einer ted Cot ae hie ol ei. Deere eel. AG aac enon 10
amereeatiniagetunidarc (hig: 20A) hex Ans rays: Salt MS pend anche p el cna Mlewha o8 eae 12

10. Gonarcus with dorso-medial projection (Fig. 12G); gonosaccus with several large gonocristae (Fig. 12C). . C. cubana
10’. Gonarcus without dorso-medial projections (Fig. 191); gonosaccus without gonocristae (Fig. 19J).......... 11

258 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

11. Apical hook of arcessus twice as long as wide (Fig. 191); apex of sternite 8 + 9 with scattered setae not in

well-defined field (Fig. 19D); forewing gradates not heavily shaded (Fig. 19B).............. C. tucumana
11'. Apical hook of arcess five times as long as wide (Fig. 18D); apex of sternite 8 + 9 with two lateral fields of long
setae (Fig. 18C); forewing gradates heavily shaded (Fig. 18B) ......-.-..--.---..--., C. tenuicornis
12. Gonarcus with dorsal plate-like projection (Fig. 20C); mesonotum pale, without spots (Fig. 18A).......... 13
12’. Gonarcus without dorsal projection (Fig. 161); paired dark spots on mesonotum (Fig. 17A) ............. 14
13. Dorsal apodeme of ectoproct prolonged ventrally as projection; a group of scales on lateral membrane of gonosaccus
adjacent to ventral projection of ectoproct (Fig. 22E); posterior apex of gonapsis smooth (Fig. 22G) . . . . C. squama
13’. Dorsal apodeme of ectoproct not prolonged ventrally as projection; no scales on lateral aspect of gonosaccus
(Fig. 20F); gonapsis with teeth at posterior apex (Fig. 20E) ..... 2... .....-...2 2000, C. dislepis
14. Dorsal apodeme of ectoproct prolonged ventrally as projection (Fig. 16C); gonosaccus with gonosetae on enormous
basal'tubercles (Fig: 161). 0.2205. 0s ene es sk 3 pee Gos oes emake an. cue d C. sanchezi
14’. Dorsal apodeme of ectoproct not forming ventral projection (Fig. 17C); gonosaccus with long setae on small basal
tibercles (Fig? 17H). %s 26.55. LF Se EE eed ta ernie GOs C. scapularis

Ceraeochrysa acmon Penny, 1998

DIAGNOSIS. — Males of this species are distinctive because of the expanded ventral tip of the
ventral fork of the dorsal apodeme of the ectoproct. The often anvil-shaped form of this expansion
gives this species a characteristic appearance and makes males of this species immediately recogniz-
able.

HEAD. — Vertex greenish yellow, unmarked. Frons and gena yellow, unmarked. Scape straw
yellow, darkened dorso-laterally (Fig. 8A); pedicel with dark band laterally; flagellum black.
Maxillary and labial palpi pale, unmarked.

THORAX. — Green. Pronotum green with dark red stripe laterally (Fig. 8A). Meso- and
metathorax unmarked. Wings: Forewing venation mostly green, except costal crossveins I—11, ori-
gin of Rs, R-Rs crossveins, inner and outer gradates and endings of anal veins dark. Length
(1.4-1.5 cm); width (0.45—0.48 cm). Hindwing venation green, except costal crossveins and gradates
dark (Fig. 8B). Hindwing length (3.25 cm): width (1.03 cm) = ratio (3.16).

ABDOMEN. — Greenish yellow with small red spots on each side of tergites. Male: Dorsal
apodeme of ectoproct with both dorsal and ventral branches widening apically (Fig. 8C). Membra-
nous wall of genital atria juxtaposed against apex of ventral branch of dorsal apodeme bearing
sclerotized plate with scales in several lines. Lateral arms of gonarcus long and triangular; medial arch
bearing medial semicircular projection with two apical horns. Arcessus elongate, thin, ending in tri-
partite apex. Gonosaccus broad, with dense field of gonocristae and long setae (Fig. 8E, F). Gonapsis
narrow and long, curved; subapically swollen; apex tapering to fine point (Fig. 8D). Apex of sternite
8 + 9 bearing lateral projections with setae having large, tubercle-like bases. Female: Subgenitale
long, with lateral lobes and median projection (cremena) (Fig. 8H). Bursa copulatrix wrinkled with
pair of large accessory glands. Spermatheca short, J-shaped (Fig. 8G).

MATERIAL EXAMINED. — Costa Rica: Limon, Amubri, A.C. Amistad, 4-21 December 1993,
G. Gallardo, holotype male (deposited in INBIO). Brazil: SAo Paulo: Jaboticabal, 20 January 1998,
Freitas, S. (221407); 10 September 1997, Freitas, S. (12); 9 December 1995, Freitas, S. (20); 23
March 1995, Freitas, S. (1 2); 22 February 1995, Freitas, S. (1 2); 12 October 1994, Gravena, R. (1 ?);
13 November 1993, Jeronimo, E. (10°); March 1999, Freitas, S. (12); Sao Simao, 2 January 1993,
Freitas, S. (1 %°)(eucalyptus); Birigui, 11 June 1994, Scomparin, C. H. J. (1°); Balsamo, 20 December
1997, Freitas, S. (1 2); Mato Grosso: Corumba, 26 July 1995, Freitas, S. (1 2); Pontes e Lacerda, 14
July 1997, Freitas, S. (1o)(rubber); Itiquira, 20 January 1994, Freitas, S. (1 2)(rubber).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 259

Ceraeochrysa caligata (Banks, 1945)

DIAGNOSIS. — This species is a member of the group of species with a recurved ventral projec-
tion of the male dorsal apodeme of the ectoproct. The pale antennae separate C. caligata from C.
claveri and C. scapularis. Ceraeochrysa caligata is probably most similar to C. cincta, but males can
be separated by the longer, more slender ventral projection of the dorsal apodeme (Figs. 9C, 10C) and
longer gonocornua (Figs. 9E, F; 10H, I) of C. caligata.

HEAD. — Green; frons and gena pale without marks. Scape and pedicel with thin dark red stripe
continued onto membranous base (Fig. 9A); flagellum pale.

THORAX. — Green; pronotum with dark red stripe laterally (Fig. 9C), mesoscutum with dark
marks (Fig. 9A); metanotum unmarked. Wing: Venation green with some transverse veins black or
darkened. Forewing: costal crossveins 4 to 10, origin of Rs, R-Rs crossveins | to 2, 4 to 8, inner and
outer gradates, Psm-Psc crossveins 3 to 5 black. Length (1.5 cm): width (0.6 cm ) = ratio (2.5).
Hindwing venation green (Fig. 9B). Hindwing length (1.42 cm): width (0.45 cm) = ratio (3.16).

ABDOMEN. — Green, unmarked. Male: Ventral arm of dorsal apodeme of tergite 9 + ectoproct
apically blunt and upturned (Fig. 9C). Gonarcus narrow, curved. Gonocornua long and decurved.
Arcessus broad, with a pair of lateral and subapical horns, apically decurved. Gonosaccus with sparse
elongate setae (Fig. 9E, F). Gonapsis thick and short (Fig. 9G). Female: Subgenitale short and thick,
forming part of a heavily sclerotized plate (Fig. 9H); bursa large sac-like; spermatheca with short ven-
tral impression (Fig. 9D).

MATERIAL EXAMINED. — Panama: Puerto Cabello, 11 June 1908, holotype male (deposited in
Museum of Comparative Zoology, Harvard University). Brazil: Sao Paulo: Jaboticabal, 24 October
1990, Silva, A. D. (1¢%)(orange); 2 September 1993, Narciso, R. (2?)(orange); January 1995,
Scomparin, C. H. J. (1o)(orange); Taquaritinga 16 October 1993, Xavier, A. L. Q. (12)(CAS); 5
March 1994, Xavier, A. L.Q. (1o%)(orange), 4 October 1993, Xavier, A. L. Q. (22)(orange);
Jaboticabal 7 April 1999, Freitas, S. (30°); Mato Grosso: Itiquira, 13 January 1997, Freitas, S.
(1o1 2)(rubber).

Ceraeochrysa cincta (Schneider, 1851)

DIAGNOSIS. — This species is a member of the group of species with a recurved ventral projec-
tion of the dorsal apodeme of the ectoproct. In Brazil, this species can be separated from C.
montoyana, C. scapularis and C. claveri by their having dark antennae. Additionally, C. scapularis
has dark spots on the mesonotum, C. montoyana has a double scape stripe and C. claveri has an api-
cally bifurcate gonapsis. Though closely related, C. caligata males have a longer, thinner ventral pro-
jection of the dorsal apodeme of the ectoproct and longer gonocornua.

HEAD. — Vertex green; gena and frons pale yellow, unmarked; maxillary and labial palpi pale;
antenna pale, apex fuscous; scape and pedicel with dorsal dark red stripe (Fig. 10A).

THORAX. — Green; pronotum with dark red stripe laterally (Fig. 10A); meso- and metanota un-
marked. Wings: Forewing venation mostly yellowish green, except costal crossveins 5—11, origin of
Rs, R-Rs crossveins 1-4, inner and outer gradates, Psm-Psc crossveins 1-4, and endings of anal veins
dark. Length (1.4 cm): width (0.5 cm) = ratio (2.96). Hindwing costal crossveins, R-Rs crossveins,
and middle part of Rs black (Fig. 10B). Length (1.11 cm): width (0.36 cm) = ratio (3.08).

ABDOMEN. — Green, unmarked. Male: Dorsal apodeme of tergite 9 + ectoproct heavily
sclerotized bearing a posteriorly-directed ventral branch (Fig. 10C). Gonarcus with vertically flat-
tened lateral arms and two caudally-directed, medial horns (gonocornua). Entoprocessus elongate,
extending well below arcessus. Arcessus with a transverse basal sclerite with two latero-apical horns,
apical sclerite with a decurved apical point. A field of tiny gonocristae arranged in rows above tergite
8 + 9 (Fig. 10H, I). Gonapsis long, slender, apically upturned (Fig. 10F). Female: Area between

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subgenitale and 9th sternite heavily sclerotized (Fig. 10D). Subgenitale with ventral invagination
without central projection (Fig. 10G). Spermatheca U-shaped, short, linked to bursa by short duct;
bursa wrinkled, sac-like (Fig. 10E).

MATERIAL EXAMINED. — Brazil: SAo Paulo: Taquaritinga, 5 March 1994, Xavier, A. L. Q.
(30°)(orange); 12 September 1992, Freitas, S. (1%)(cotton); 20 September1993, Xavier, A. L. Q.
(22); 12 September 1992, Freitas, S. (5); Jaboticabal, 9 December 1995, Freitas, S. (102); 30 Sep-
tember 1997, Freitas, S. (1 2)(CAS); 2 October 1997, S. Freitas (1 o")(CAS); Mato Grosso: Itiquira, 20
January 1997, Freitas, S. (1%); 14 June 1997, Freitas, S. (1 ?)(rubber).

Ceraeochrysa claveri (Navas, 1911)

DIAGNOSIS. — This species is often more yellow in coloration than others of the genus. It is a
member of the group of species with males having a recurved ventral projection of the dorsal apodeme
of the ectoproct. This is one of three Brazilian species with dark antennae. However, C. montoyana
has a double dorsal scape stripe, and C. scapularis has dark spots on the mesonotum. Males of C.
claveri also have a distinctive thick apical fork of the gonapsis.

HEAD. — Vertex golden yellow; frons and gena pale, unmarked; pedicel and scape with a dorsal
mid-line red stripe continued onto the antennal basal membrane as a triangular spot; flagellum black
(Fig. 11A); maxillary and labial palpi pale.

THORAX. — Green; pronotum yellow with dark red stripe laterally (Fig. 11A); meso- and
metanota unmarked. Wings: Venation mostly green with some crossveins black: forewing costal
crossveins 4—12, origin of Rs base, R-Rs crossveins, Rs-Psm crossveins 1—4, gradates, Psm-Psc
crossveins 6-8, unforked marginal veins, and cubital and anal vein endings dark. Length (1.4 cm):
width (0.5 cm) = ratio (3.08). Hindwing costal crossveins 5—13, R-Rs crossveins, and gradates dark
(Fig. 11B). Length (1.37 cm): width (0.42 cm) = ratio (3.26).

ABDOMEN. — Green, unmarked. Male: Dorsal apodeme of tergite 9 + ectoproct bearing ventral
fork which is apically acute (Fig. 11C). Gonapsis long, slender, posteriorly forked (Fig. 11F).
Gonarcus not arched. Entoprocessus long, caudally pointed. Arcessus short, membranous, triangular
in dorsal view, without dorsal horns. Gonosaccus with sparse gonosetae (Fig. | 1H, I); above apex of
9th sternite a field of gonocristae (Fig. 11E). Female: Area between subgenitale and 7th sternite
heavily scletotized (Fig.1 1D). Bursa copulatrix with a pair of large glands (Fig. 11G). Spermatheca
elongate, U-shaped, ventral impression elongate.

MATERIAL EXAMINED. — Brazil: Sao Paulo: Taquaritinga, 16 October 1993, (322); 5 March
1994, Xavier, A. L. Q. (20°22)(orange); 7 January 1993, Freitas, S. (2%)(corn); 30 March 1993,
Freitas, S. (2o°12)(guava); Mato Grosso: Itiquira, 7 September 1999, Freitas, S. (30°42 )(rub-
ber)(SDF/CAS).

Ceraeochrysa cubana (Hagen, 1861)

DIAGNOSIS. — This is a rather small species with pale antennae and considerable dark markings
on the forewings. Females are indistinguishable from the more northern C. valida. Males have a num-
ber of autapomorphies: abdominal spiracles are unusually enlarged; the apex of sternite 8 + 9 is bifur-
cate and spiny; there are several large gonocristae on the gonosaccus; and the gonarcus has a vertical
dorso-medial plate. Males also have a rugose vertex which is otherwise only found in C. paraguaria
and C. scapularis.

HEAD. — Green; vertex glossy on female and rugose on male, unmarked; scape with a dark red,
thin stripe dorso-laterally (Fig. 12A); flagellum pale; frons and gena unmarked.

THORAX, — Green. Pronotum with light red stripes laterally (Fig. 12A). Meso- and metanota un-
markec. VYings: Venation green with black crossveins. Forewing costal crossveins 2—15, origin of Rs,

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 261

R-Rs crossveins |—8, inner and outer gradates, Psm-Psc crossveins 3-8, and cubital and anal veins
apical endings dark. Length (1.15 cm): width (0.37 cm) = ratio (2.97). Hindwing costal crossveins
4-12, R-Rs veins 1-4, and inner and outer gradates dark (Fig. 12). Length (1.06 cm): width (0.32 cm)
= ratio (3.31).

ABDOMEN. — Green, unmarked. Male: Dorsal surface of 9th tergite + ectoproct deeply grooved;
apex of 9th sternite with pair of lateral projecting forcipate lobes (Fig. 12D); dorso-laterally to ninth
sternites two lobes bearing teeth and setae (Fig. 12C). Gonarcus broad with a dorso-medial, plate-like
projection. Entoprocessus short. Arcessus large, apico-medially, decurved and acute. Gonosaccus
with long setae (Fig. 12F, G). Gonapsis long and wide. Female: Spermatheca short. Bursa sac-like,
wrinkled (Fig. 12E).

MATERIAL EXAMINED. — Cuba: Soledad, near Cienfuegos, 6-20 August, Banks, N., neotype
male (deposited in Museum of Comparative Zoology, Harvard University), Brazil: S40 Paulo:
Taiuva, 23 July 1990, Scatolin, M. (22)(orange); 23 October 1990, Scatolin, M. (30)(orange);
Taquaritinga, 8 January 1993, Xavier, A. L. Q. (3@)(orange); 25 March 1993, Xavier, A. L. Q. (52);
Jaboticabal, 22 February 1995, Freitas, S. (400%352)(corn, guava, cotton, eucalyptus)(SDF/CAS);
Mato Grosso: Itiquira, 20 January 1997, Freitas, S. (3%)(rubber).

Ceraeochrysa everes (Banks, 1920)

DIAGNOSIS. — This species is a member of the everes species group with males having
thick-based spines at the apex of sternite 8 + 9, a broad dorsal plate medially on the gonarcus; broad,
vertically oriented entoprocesses; vertically elongate lateral arms of the gonarcus, and a field of
gonocristae laterally below the apex of the ectoproct. Of these species, the more northern C.
costaricensis has two dark spots on the pronotum rather than longitudinal stripes. Ceraeochrysa
everes has a field of gonosetae which line up in rows and evenly curved ventral margin of the
ectoproct, while C. squama has a randomly arranged field of gonosetae and a sclerotized ventrally
projecting arm of the ectoproct at the apex of the dorsal apodeme.

HEAD. — Greenish yellow; frons and gena unmarked; maxillary and labial palpi pale; scape and
pedicel green with middle of lateral surface bearing reddish brown diffuse spot, sometimes poorly de-
fined; flagellum black (Fig. 13A).

THORAX. — Green. Pronotum somewhat yellowish with lateral reddish to chocolate brown
stripe (Fig. 13A). Meso- and metanota unmarked. Wings: Venation mostly green, except for some
crossveins dark (Fig 13B): Forewing costal crossveins in middle part, R-Rs crossveins at apical ends,
and inner and outer gradates dark. Length (1.52 cm): width (0.46 cm) = ratio (3.0) Hindwing inner
gradates dark. Length (1.35 cm): width (0.43 cm) = ratio (3.14).

ABDOMEN. — Green, unmarked; Male: Tergite 9 + ectoproct without visible ventral projection
of apodeme (Fig. 131). Gonapsis flattened, apex plate-like and upturned with row of small apical teeth
and one larger dorso-apical tooth (Fig. 13H). Gonarcus strongly arched horizontally with a vertically
oriented, medio-dorsal plate; lateral arms crescent-shaped. Arcessus membranous basally with a pair
of hooks and two inflated lobes; apex bearing a medial hook and pair of lateral, decurved and medi-
ally-curved sclerotized lobes. Gonosaccus large with dense medial field of stout setae; two lateral
lobes of gonosaccus with long setae, and between medial field of short gonosetae and lateral field of
long gonosetae a field of tiny gonocristae. Lateral to the genital atria a field of tiny scales arranged in
several lines. Apex of sternite 8 + 9 with large basal tubercles bearing setae arranged on two lateral
lobes (Fig. 13C, D, E). Female: Spermatheca elongate, twisted, opening into a transversally wrinkled
bursa (Fig. 13G). Subgenitale heavily sclerotized with a median projection (cremena), attached to
bursa by a sclerotized plate.

MATERIAL EXAMINED. — French Guiana: Roches de Kourou, July, Le Moult, lectotype female
(deposited in Museum of Comparative Zoology, Harvard University), Brazil: Sao Paulo:

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Jaboticabal, 20 January 1998, Freitas, S. (4%)(corn)(SDF/CAS); 12 February 1995, (1o)(guava);
Luiz Antonio, 8 December 1993, Freitas, S. (1o*)(eucalyptus); Taquaritinga, 20 November 1992,
Freitas, S. (1%)(orange); 8 January 1993, Xavier, A. L. Q. (1 ?)(orange); 5 March 1994, Xavier,
A. L. Q. (1ce)(orange); Balsamo, 20 December 1997, Freitas, S. (1 o)(rubber); Mato Grosso: Itiquira,
10 September 1996, Scomparin, C. H. J. (2¢)(rubber),; 23 September 1996, Scomparin, C. H. J.
(12)(CAS).

Ceraeochrysa montoyana (Navas, 1913a)

DIAGNOSIS. — This species is closely related to other cincta group species that have dark anten-
nae, such as C. claveri. However, in C. claveri the scape has a single stripe that continues onto the
antennal basal membrane, and the spermatheca of C. claveri is not U-shaped. In addition, the
two-striped scape of C. montoyana is a distinctive characteristic.

HEAD. — Vertex golden yellow; gena and frons pale yellow; scape yellow, with two red stripes,
one dorso-laterally and the other dorso-centrally, the central one continuing onto antennal basal mem-
brane (Fig. 14A); pedicel with red markings; flagellum black.

THORAX. — Green. Pronotum green with lateral dark red stripes (Fig. 14A). Meso- and metanota
green, unmarked. Wings: Forewing venation mostly green, except costal crossveins, R-Rs
crossveins, first Rs-Psm crossveins, Psm-Psc, apical and anal endings dark. Length (1.5 cm): width
(0.54 cm) = ratio (2.78). Hindwing venation mostly green, except R-Rs crossveins dark (Fig. 14B).
Length (1.38 cm): width (0.45 cm) = ratio (3.07).

ABDOMEN. — Green, without marks. Female: Spermatheca V-shaped, short, with long
spermathecal gland (Fig. 14C). Subgenitale with two anterior lobes (Fig. 14D).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 25 March 1999, Freitas, S.
(1 2)( guava).

Ceraeochrysa paraguaria (Navas, 1920) new combination

DIAGNOSIS. — This species is easily recognized by the four round small spots on the pronotum.
Sometimes C. paraguaria is confused with Plesiochrysa brasiliensis when only external morphologi-
cal characteristics are considered. However, P. brasiliensis is a larger, more robust species, with
pronotum that is longer than wide. The pronotal spots of C. paraguaria are much darker and
well-delimited, not diffuse. Finally, the bifurcate apex of sternite 8 + 9 in males is found only in C.
cubana and C. scapularis.

HEAD. — Green; frons and gena unmarked; scape, pedicel and flagellum pale, unmarked.

THORAX. — Green. Pronotum with pair of dark red small spots laterally on either side (Fig. 15A).
Meso- and metanota unmarked. Wings: Venation green, except for certain crossveins: forewing
costal crossveins 4-10, origin of Rs, R-Rs crossveins, Psm-Psc crossveins |—3, cubital and anal veins,
imc lateral veins, and inner and outer gradates dark. Length (1.25 cm): width (0.41 cm) = ratio (3.04).
Hindwing costal crossveins dark (Fig. 15B). Length (1.08 cm): width (0.33 cm) = ratio (3.27).

ABDOMEN. — Green, unmarked. Male: Tergite 9 + ectoproct deeply grooved dorsally forming
two separate lobes. Sternite 8 + 9 apically bifurcate. Gonarcus narrow, lateral arms projecting verti-
cally. Entoprocessus short and wide. Gonocornua long, tapered, reaching apex of arcessus. Arcessus
triangular; apically acutely decurved. Gonosaccus with many long setae (Fig. 15H, I). Gonapsis slen-
der with multi-pronged apical appendage (Fig. 15E). Female: Subgenitale short and wide (Fig. 15F).
Spermatheca short, with thick, twice-coiled spermathecal gland.

MATERIAL EXAMINED. — Brazil: Sao Paulo: Taquaritinga, 7 January 1993, Freitas, S.
(90°12 (orange); Jaboticabal, 17 February 1995, Freitas, S. (120)(orange); 20 October 1994, Cha-

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 263

gas, A. (2?)(orange); 7 August 1996 Freitas, S. (40°42) (guava); 20 October 1996, Freitas, S.
(101 2)(CAS).

REMARKS. — Navas described Chrysopa paraguaria in 1920. Later, Navas (1924) modified the
name to C. paraquaria. However, Article 32.5.1 of the International Code of Zoological Nomencla-
ture (ICZN 1999) states that “If there is in the original publication itself, without recourse to any exter-
nal souce of information, clear evidence of an inadvertent error, such as a lapsus calami or a copyist’s
or printer’s error, it must be corrected. Incorrect transliteration or latinization, or use of an inappropri-
ate connecting vowel, are not to be considered inadvertent errors.” Thus, without any indication in the
original publication that this name was incorrectly spelled we have maintained the original spelling.
Brooks and Barnard (1990) listed this species in Plesiochrysa, but the male genitalia, including elon-
gate gonapsis, clearly indicate placement of this species in Ceraeochrysa.

Ceraeochrysa sanchezi (Navas, 1924b)

DIAGNOSIS. — This is one of the species with dark antennae which can be separated most easily
by characteristics of the male genitalia. The elongate ventral projection of the ectoproct and subapical
dorsal projection of the ninth sternite are characteristic of this species, as is the pair of triangular fields
of gonocristae projecting above and medial to the apex of the ninth sternite.

HEAD. — Yellow; frons and gena unmarked; maxillary and labial palpi pale; scape and pedicel
yellow with lateral stripe brown, continued onto antennal basal membrane (Fig. 16A). Flagellum
black. Maxillary and labial palpi pale.

THORAX. — Yellow. Pronotum with lateral stripe thick, red. Mesopraescutum and mesoscutum
with lateral red spots (Fig. 16A). Wings: Venation mostly green with the following transverse veins
black: forewing costal crossveins, origin of Rs, R-Rs crossveins; inner and outer gradates, Psm-Psc
crossveins, anal and cubital vein apices, marginal forks and unforked apices. Length (1.35 cm): width
(0.45 cm) = ratio (2.96). Hindwing veins: costal crossveins 4—10; R-Rs crossveins 5—8; inner and
outer gradates dark (Fig. 16B). Length (1.14 cm): width (0.35 cm) = ratio (3.26).

ABDOMEN. — Yellow, without dark marks. Male: Tergite 9 + ectoproct with finger-like dorsal
projection of dorsal apodeme above callus cerci and elongate ventral lobe of dorsal apodeme project-
ing ventrally from ectoproct (Fig. 16C). Sternite 8 + 9 with lateral concavities, which receive ventral
ectoproct projection and subapical dorsally-projecting lobes (Fig. 16D). Gonapsis slender, apex with
multi-lobed, spiny appendages (Fig. 16G). Gonarcus thick, lateral arms triangular in shape.
Entoprocessus quite broad, apically obtusely pointed. Arcessus triangular, apex forming medial
decurved point. Gonosaccus with three pair of long setae originating from stout bases. Posterior to
gonosaccus a pair of triangular lobes with fields of gonocristae (Fig. 161, J). Female: Bursa wrinkled,
with a pair of large accessory glands (Fig. 16H). Spermatheca U-shaped with distal portion dilated
(Fig. 16F). Subgenitale with median projection (cremena) and basal sclerotized plate (Fig. 16E).

MATERIAL EXAMINED. — Cuba: Havana, Playa Chivo, 18 December 1923, Cervera, holotype
male (deposited in Museum of Comparative Zoology, Harvard University), Brazil: Rio Grande do
Norte: Assu, 6 June 1991, Freitas, S. (20°2?)(cantaloupe); Sao Paulo: Jaboticabal, 5 March 1992,
Freitas, S. (1 2)(orange); 26 January 1995, Freitas, S. (1 2 1.o)(guava); 2 May 1995, Seguin, L. D. (1%);
4 December 1996, Freitas, S. (12)(CAS); 10 February 1998, Freitas, S. (1o°)(CAS); Taquaritinga, 7
January 1993, Freitas, S. (2°)(orange); Mato Grosso, Itiquira, 7 September 1999, Freitas, S.(20l ?).

Ceraeochrysa scapularis (Navas, 1914)
DIAGNOSIS. — This is a small species most closely related to C. cubana and C. paraguaria. All

three species have a bifurcate apex of sternite 8 + 9 and gonosetae with large bases. Of the three, C.
scapularis is the only one with dark antennae and two dark spots on the mesonotum.

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HEAD. — Yellowish green. Gena and frons unmarked. Scape with lateral dark red stripe, pedicel
dark-ringed; flagellum black (Fig. 17A). Maxillary and labial palpi pale.

THORAX. — Greenish yellow. Pronotum as long as wide with lateral red stripe (Fig. 17A).
Mesonotum with pair of small dark red spots (Fig. 17A). Wings: Green longitudinal veins and black
crossveins. Forewing crossveins darker than hindwing veins: costal crossveins |—15, origin of Rs;
R-Rs crossveins 1—9; inner and outer gradates; Psm-Psc crossveins |—7; anals, cubitals and unforked
marginal endings black (Fig. 17B). Length (1.1 cm): (0.35 cm) = ratio (3.03). Hindwing costal
crossveins; R-RS crossveins, Rs-Psm crossveins, inner and outer gradates, and bases of Psm-Psc
crossveins black. Length (0.99 cm): width (0.29 cm) = ratio (3.41).

ABDOMEN. — Greenish yellow. Male: Tergite 9 + ectoproct deeply grooved dorsally. Apex of
sternite 8 + 9 with two forcipate lobes (Fig. 17D). Gonarcus thick. Gonocornua slender and upturned.
Arcessus triangle-shaped, apex tapered into a medial point. Gonosaccus with apical field of long setae
(Fig. 17H, I). Posterior to gonosaccus two large lateral lobes with strong gonocristae and medial field
of small gonocristae (Fig. 17C). Gonapsis elongate, narrow, widening subapically (Fig.17E). Fe-
male: Subgenitale without median projection (Fig. 17G). Bursa with two large accessory glands (Fig.
17F). Spermatheca U-shaped with broad spermathecal duct.

MATERIAL EXAMINED. — Brazil: SAo Paulo: Catanduva, January 1995, Silva, J. L. (1@)(or-
ange); Luiz Antonio, 9 December 1991, Freitas, S. (1o)(eucalyptus); 5 October 1993, Crispolin, F.
(32 )(eucalyptus); Jaboticabal, 17 February 1995, Freitas, S. (10c)(orange); 23 March 1995, Freitas,
S. (1o%)(orange); 14 July 1995, Freitas, S. (1 “)(orange); 17 February 1995, Freitas, S. (4? )(orange); 7
August 1996, Freitas, S. (20°)(CAS).

Ceraeochrysa tenuicornis Adams and Penny, 1987

DIAGNOSIS. — This species cannot be separated with certainty from other species of
Ceraeochrysa without study of the male genitalia. Of the male genital elements, the elongate
gonocornua and ventral field of small gonosetae on the gonosaccus are most characteristic.

HEAD. — Yellowish green. Frons, gena and vertex unmarked. Scape, pedicel and flagellum pale;
scape with a tiny dark spot on apico-lateral margin (Fig. 18A).

THORAX. — Pale green. Pronotum yellow with pale red stripe laterally (Fig. 18A). Meso- and
metanota green, unmarked. Wings: Venation green, except some costal crossveins, R-Rs crossveins,
gradates darkened; gradates darkly bordered. Forewing stigma unmarked. Length (0.96 cm): width
(0.33 cm) = ratio (2.91). Hindwing venation green (Fig. 18B). Length (0.83 cm): width (0.25 cm) =
ratio (3.32).

ABDOMEN. — Green. Tergites large, unmarked. Male: Dorsal apodeme of tergite 9 + ectoproct
dark, forked at callus cerci, which has a few trichobothria (Fig. 18C). Sternite 8 + 9 densely covered
by setae on conspicuous tubercles. Gonarcus thick, hardly arched, with fan-like lateral arms.
Entoprocessus short and sharply decurved. Gonocornua straight, angled upward, reaching almost 2/3
length of arcessus. Arcessus divided: basal sclerite bearing two small horns and apical narrow, slender
sclerite terminally decurved. Gonosaccus bearing laterally two pockets of long setae and ventrally a
lobe with large field of tiny gonosetae, between them some small points ina vertical line (Fig. 18D, E).
Gonapsis straight, unforked (Fig. 1 8F).

MATERIAL EXAMINED. — Brazil: Amazonas, Manaus, INPA campus, 23 April 1976, I. S.
Goreyeb, holotype male (deposited at INPA), Mato Grosso: Itiquira, 20 January 1997, Freitas, S.
(1 o)(rubber).

REMARKS. — This is a species not known from south of the Amazon Basin and no females were
collected during this project. For a description of the female, see Adams and Penny (1987).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 265

Ceraeochrysa tucumana (Navas, 1919), new status

DIAGNOSIS. — Ceraeochrysa tucumana is a small species that resembles Chrysoperla externa in
wing venation, but has stripes on the antennal scape and pronotum. The male genitalia and sternite
8 + 9 resemble C. /ineaticornis (C. A. Tauber et al. 2000) and C. berlandi (Museum of Comparative
Zoology material), but they have dark antennae and do not have the field of gonocristae below the
gonosaccus.

HEAD. — Greenish yellow. Frons and gena without marks. Maxillary and labial palpi pale. Scape
with lateral brown stripe, pedicel dark-ringed; flagellum pale (Fig. 19A).

THORAX. — Pale green. Pronotum with lateral stripe reddish brown (Fig. 19A). Meso- and
metanota unmarked. Wings: Venation mostly green with some transverse veins black: forewing
costal crossveins 1—13, origin of Rs, Rs-Psm crossveins |—3, inner and outer gradates, and Psm-Psc
crossveins. Length (1.09 cm): width (0.37 cm) = ratio (2.96). Hindwing costal crossveins I-13, R-Rs
crossveins; and inner and outer gradates dark (Fig. 19B). Length (0.92 cm): width (0.30 cm) = ratio
(3.07).

ABDOMEN. — Greenish yellow with pale red spots on the tergites 2 and 3. Male: tergite 9 +
ectoprocts deeply divided dorsally making two lateral lobes (Fig. 19E); apex of sternite 8 + 9 with
pointed projection and thick-based setae (Fig. 19D). Gonarcus thick. Entoprocessus elongate, thin,
decurved. Gonocornua vertically curved. Arcessus large, with medio-dorsal forked horn; apex up-
turned as sclerotized lobe with apical hook (Fig. 191, J). Gonapsis slender (Fig. 19H). Gonosaccus
with two lateral, protruding fields of long setae. Between gonosaccus and sternite 8 + 9 a membra-
nous area with gonocristae. Female: Spermatheca short with thick spermathecal gland (Fig. 19F).
Subgenitale with short median projection (Fig. 19G).

MATERIAL EXAMINED. — Brazil: SAo Paulo: Taquaritinga, 8 January 1993, Xavier, A. L.
(1o)(orange); 1 June 1994, Narciso, R. (1 2)(orange); Luiz Antonio, 8 September 1993, Freitas, S.
(1¢)(eucalyptus); 21 September 1993, Freitas, S. (3o°)(eucalyptus); 8 November 1993, Freitas, S.
(201 2)(eucalyptus); 21 December 1993, Freitas, S. (573 2)(eucalyptus); 5 October 1993, Crispolin,
F. (32 )(eucalyptus); 13 July 1992, Freitas, S. (1 2)(eucalyptus); 29 April 1994, Freitas, S. (1 ?)(CAS);
8 December 1998, Freitas, S. (1 2)(CAS).

REMARKS. — In his original description, Navas indicated that the type (sex not specified) was de-
posited in the museum at La Plata, Argentina. In their world list of chrysopid species, Brooks and
Barnard (1990) placed this species as an “incertae sedis” at the end of the list. Phillip Adams found the
type male at the Buenos Aires Museum, cleared and stained the genitalia, and made detailed sketches.
These sketches were used to identify the above material. The genital elements include an elongate
gonapsis, clearly placing this species in Ceraeochrysa.

Ceraeochrysa dislepis de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Bra-MT-Itiquira, P. E. Michelin 20/1/97, Freitas, S. Leg. 1407."

DIAGNOSIS. — This species is a member of the everes species group with males having
thick-based setae at the apex of sternite 8 + 9; a thick medial field of small gonosetae on the
gonosaccus; a broad dorsal plate on the gonarcus; elongate, vertically-tapering, lateral arms to the
gonarcus; and a pair of distinctive dorsal projections just posterior to the gonarcus. Of the four known
species in the group, two [C. costaricensis (Penny 1997) and C. squama] have ventral projections
from the ectoproct and gonapsis with lateral wings, while the other two species (C. everes and C.
dislepis) have no ventral projection from the ectoproct and a gonapsis terminating in a vertical plate
with teeth. Although similar, C. dis/epis can be separated from C. everes by the central indentation of

266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

the medial plate of the gonarcus and the lack of a field of tiny gonocristae laterally on the membrane
below the ectoproct.

The name comes from “dis” meaning separated and “lepis” scales, describing the two lateral
fields of small, scaly gonocristae on the gonosaccus.

HEAD. — Yellowish green. Vertex, frons and gena unmarked. Maxillary and labial palpi pale
yellow, unmarked. Scape yellow with lateral brown stripe widening apically; pedicel as scape;
flagellum black (Fig. 20A).

THORAX. — Green. Pronotum green with a dark red lateral stripe (Fig. 20A). Meso- and
metanota unmarked. Wings: Forewing longitudinal veins green; costal crossveins, R-Rs crossveins,
inner and outer gradates, anterior intersection of Psm-Psc crossveins, forked and unforked marginal
veins, and margins near forks darkened. Length (1.3 cm): width (0.4 cm) = ratio (3.25). Hindwing
apically acutely pointed, green except costal crossveins; R-Rs crossveins, apex of Rs, and outer
gradates dark (Fig. 20B). Length (1.14 cm): width (0.37 cm) = ratio (3.08).

ABDOMEN. — Green, unmarked. Male: Dorsal apodeme of tergite 9 + ectoproct without ventral
projection. Gonapsis flattened, anterior apex a vertical plate bearing two rows of teeth (Fig. 20E).
Gonarcus lightly curved with dorsal medially-indented plate (Fig. 20C). Arcessus with dorso-lateral
inflated lobes, a pair of small dorsal acute lobes, and decurved apical median hook with lateral lobes
(Fig. 20D). Gonosaccus with lateral field of long setae and ventral fields of tiny gonocristae below
setal fields (Fig. 20D). Apex of sternite 8 + 9 bearing setae with thick bases arranged on small lateral
lobes (Fig. 20F). Female: Spermatheca twisted, but not coiled, apex with lateral alate projection (Fig.
20G). Bursa a large, transversely wrinkled sac with pair of large accessory glands.

OTHER MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 11 January1997, Freitas, S.
(12 paratype)(rubber); 20 January 1997, Freitas, S. (1% paratype)(rubber); 14 September1998,
Freitas, S. (1c paratype)(rubber); 10 September 1996, Scomparin, C. H. J. (1 ? paratype)(rubber); 4
November1996, Scomparin, C. H. J. (221o paratypes)(rubber)(SDF/CAS); 30 December 1996,
Scomparin, C. H. J. (1o*)(rubber); 18 July 1998, Freitas, S. (1? paratype)(rubber); 18 November
1996, Scomparin, C. H. J. (lo paratype)(rubber); S40 Paulo: Jaboticabal, 22 November 1995,
Freitas, S. (12 paratype)(orange); 7 July 1996, Freitas, S. (10 paratype)(CAS); 30 September 1997,
Freitas, S. (1% paratype)(orange); 7 July 1998, Freitas, S. (1o paratype)(orange); 17 November 1995,
Freitas, S. (1 2 paratype)(orange); November 1995, Freitas, S. (1 * paratype)(orange); Luiz Antonio, 8
December 1993, Freitas, S. (1c paratype)(eucalyptus); 21 December 1993, Freitas, S. (Ie
paratype)(eucalyptus); Goias: Cachoeira Alta, 10 June 1997, Freitas, S. (1 2 paratype)(rubber).

Ceraeochrysa dolichosvela de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled, “Bra-SP-Jaboticabal, FCAV, 20/1/98, 69, Freitas, S. Leg.”

DIAGNOSIS. — The long, uncoiled shape of the spermatheca is not found in any other species.

The name “dolichovela” comes from the Greek dolichos meaning long, and refers to the long
spermatheca and its vela in this species.

HEAD. — Vertex, gena and frons greenish yellow. Unmarked. Scape yellow-orange; pedicel
black; flagellum black. Maxillary and labial palpi pale (Fig. 21 A).

THORAX. — Green with a pale stripe on mid-line. Pronotum green with pale brown-red lateral
stripe (diffuse in paratype) (Fig. 21A). Meso- and metanota unmarked. Wings: Venation mostly
green, except costal crossveins, R-Rs transverse veins and gradates black. Length (1.49 cm): width
(0.5 cm) = ratio (2.88). Hindwing venation mostly green, except R-Rs transverse veins and outer
gradates black (Fig. 21B). Length (1.26 cm): width (0.41 cm) = ratio (3.07).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 267

ABDOMEN. — Green with pale stripe along mid-line. Female: Spermatheca extremely long,
slender, uncoiled; with long spemathecal duct (Fig. 21C). Posterior part of subgenitale bilobed, with
large medial anterior projection.

OTHER MATERIAL EXAMINED. — Brazil: Mato Grosso do Sul: Corumba, 23 July 1995, Freitas,
S. (12 paratype)(orange).

Ceraeochrysa squama de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP, 20/01/95, SP14A, Freitas, S.” (guava).

DIAGNOSIS. — This species is a member of the everes species group with character states listed
under C. dis/epis. The lateral arms of the gonapsis relate it most closely with C. costaricensis Penny,
1997, but it differs in having dark antennae and red stripes rather than spots laterally on the pronotum.

The name “squama” comes from the Latin for scale, and refers to the two fields of tiny
gonocristae laterally on gonosaccus.

HEAD. — Vertex, frons and gena golden yellow, unmarked. Maxillary and labial palpi pale.
Scape golden yellow, with small brown spot at apical dorso-lateral corner; pedicel with lateral brown
stripe; flagellum black (Fig. 22A).

THORAX. — Green. Pronotum green with lateral dark red stripe. Meso- and metanota unmarked
(Fig. 22A). Wings: Forewing venation mostly green with some transverse veins darkened, such as
costal crossveins and R-Rs crossveins. Length (1.3 cm): width (0.46 cm) = ratio (2.83). Hindwing ve-
nation mostly green, except inner and outer gradates dark (Fig. 22B). Length (1.17 cm): width
(0.36 cm) = ratio (3.25).

ABDOMEN. — Yellowish green, unmarked. Male: Dorsal apodeme of tergite 9 + ectoproct
strongly sclerotized and with ventral projection (Fig. 22E). Gonapsis flattened, apex turned up as ver-
tical plate-like lobe, but without teeth; subapical part with lateral alate projection apically decurved
(Fig. 22G). Gonarcus strongly arched, with median plate-like projection as long as wide; lateral arms
broadly rounded ventrally. Two inflated dorsal projections posteriad of gonarcus. Arcessus membra-
nous basally with pair of small diverging lobes; apex medially acute and lateral membranous lobes.
Gonosaccus large, with dense central field of small, stout setae; two lateral lobes with long setae
flanked by a field of tiny gonocristae. Plate of randomly organized, sclerotized, pointed scales later-
ally on membranous wall of genital atrium, probably juxtaposed with ventral projection of ectoproct
when depressed (Fig. 22A, B, C).

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 30 September 1997, Freitas,
S. (1¢% paratype); 24 April 1999, Takahashi, K. (1% paratype)(guava).

Genus Chrysoperla Steinmann, 1964

This is probably the most studied genus in the family. Adults are abundant in north temperate
fields and yards, and are attracted to lights. They feed on grass pollen and are the most frequently con-
sidered candidates for inundative releases in fields and orchards for biological control. They are being
mass-reared commercially for biological control programs in the U.S.A. (Tauber et al. 2000).

In a recent revision of the genus, Brooks (1994) found 35 species distributed world-wide, but
with a preponderance of species in the Holarctic Region. Adult males of this genus have one distinc-
tive structural feature. Sternite 8 + 9 of males is subapically constricted, giving the apex the appear-
ance of asmall apical lobe (Figs. 23E, 24D, 25C). All New World species have red genae, and lateral
setae on the pronotum have thickened bases. However, other chrysopids have red genae and females
must be identified with great caution. In this study we have found three species; the adults can be dis-
tinguished using the following key.

268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

KEY TO THE SPECIES OF CHRYSOPERLA IN BRAZILIAN AGRO-ECOSYSTEMS
(modified from Brooks, 1994)

IeForewinpieradates greens 2-2. 2. iy 4 c'o ska 3 ES Nee 1 eee ee y)
le Forewing eradatesi blacks... fae. 0 © 2 Fs tase ei chee See es 2 C. defreitasi
2. Male arcessus only weakly decurved apically (Fig. 241)... 2.2... 6... eee eee ee ee eee C. externa
2 Male arcessus strongly decurved apically (Fig: 25E))<) <<... .-igs a9? Se ee Ae eee C. raimundoi

Chrysoperla defreitasi Brooks, 1994

DIAGNOSIS. — Chrysoperla defreitasi is closely related to C. externa and C. raimundoi.
Chrysoperla defreitasi may be differentiated from the other two species by black gradate crossveins
vs. green crossveins in C. externa and C. raimundoi.

HEAD. — Greenish yellow; red gena, lateral clypeus and lateral labrum; narrow red stripe on
fronto-clypeal suture; antenna with scape and pedicel pale yellow, flagellum pale; maxillary palpi
black, marked dorsally (Fig. 23C).

THORAX. — Yellowish green with yellow stripe on dorsal mid-line; Pronotum with narrow red-
dish lateral stripe (Fig. 23A); meso- and metanota unmarked. Wings: Forewing venation mostly
green with black gradates, cubital and anal veins black; forewing length (1.3 cm): width (0.37cm) =
ratio (3.5). Hindwing length (1.19 cm): width (0.37 cm) = ratio (3.22).

ABDOMEN. — Green with yellow stripe on mid-line. Male: long apodeme of tergite 9 forked on
callus cerci; lip of sternite 8 + 9 short (Fig. 23E); many gonocristae; gonarcus large, arched; arcessus
broad, tapered subapically, apex curved; gonosaccus with many gonosetae (Fig. 23F, G); tignum
elongate, acumen long (Fig. 23D).

MATERIAL EXAMINED. — Brazil: Jaquaritingas, 15 November 1992, S. de Freitas, holotype
male (MZ, USP), Sao Paulo: Jaboticabal, 7 August 1995, Seguim, L. D. (1%); August 1996, Freitas,
S. (12); 16 December 1998, Freitas, S. (10); Mato Grosso: Itiquira: 7 September 1999, Freitas, S.
(502 2)(rubber)(SDF/CAS).

Chrysoperla externa (Hagen, 1861)

DIAGNOSIS. — This species can be separated from C. defreitasi by the dark gradate veins of the
forewing in C. defreitasi, and from C. raimundoi by the male’s more exaggerated curvature of the tip
of the arcessus in C. raimundoi.

HEAD. — Vertex, greenish yellow, flattened with several transverse striations (Fig. 24A); frons
and clypeus unmarked; gena with red spot not reaching ventral margin near maxillae; black dorsal
marks on maxillary palpi (Fig. 24C); scape and pedicel pale yellow; flagellum basally pale and api-
cally fuscous.

THORAX. — Green with yellow stripe on dorsal mid-line; pronotum with grayish latero-anterior
spot; base of setae black (Fig. 24A); meso- and metanota without marks. Wings: Venation yellowish
green (Fig. 24B); forewing length (1.23 cm): width (0.31 cm) = ratio (4.0). Hindwing length
(1.19 cm): width (0.36 cm) = ratio (3.31).

ABDOMEN. — Greenish yellow with yellow stripe on dorsal mid-line, unmarked. Male: Tignum
elongate, narrow with medial acumen small, triangular (Fig. 24E); gonarcus hardly curved; arcessus
narrow, apex slightly curved; gonosaccus with many long gonosetae (Fig. 241, J); numerous
gonocristae (Fig. 24H). Female: Long subgenitale (Fig. 24G); spermatheca short (Fig. 24F).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Taiuva, 19 July 1990, Pazini, W. (60°62 )(orange);
Jaboticabal, 10 August 1992, Freitas, S. (20°)(guava); Taquaritinga, 7 January 1993 Freitas, S.
(601 $ (orange).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 269

REMARKS. — Chrysoperla externa is probably the most common chrysopid in the Neotropical
region and occurs associated with many graminous plants, apple trees, cotton, cantaloupe, eucalyptus,
rubber and cashew nut.

Chrysoperla raimundoi de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP, FCAV, 20-04-95, 1’, Freitas, S.”

DIAGNOSIS. — Chrysoperla raimundoi closely resembles C. externa and C. defreitasi. From C.
externa it can be differentiated on cephalic characters: the vertex of C. externa is as long as wide and
less striated. The head is also conspicuously different between them: C. raimundoi has a more elon-
gate clypeus. The male genitalia also display some differences, e.g., the apex of the arcessus of C.
raimundoi is quite curved and the gonosaccus bears longer gonosetae. Chrysoperla defreitasi can be
differentiated from C. raimundoi by the smooth vertex, frons and red clypeus as well as the dark
gradate veins of the forewing of C. defreitasi.

The name “raimundoi” is given in honor of Agricultural Engineer Raimundo José Ferreira who
collected the type series.

HEAD. — Pale green; vertex raised with several transverse striations; diffuse red post-ocular
spot; frons yellowish without marks, gena red throughout; dorsally black maxillary and labial palpi;
scape and pedicel without marks (Fig. 25A).

THORAX. — Green with thick median yellow band; pronotum with lateral narrow reddish brown
band not reaching the posterior corner. Wings: Venation green (Fig. 25B); forewing length (1.22 cm):
width (0.40 cm) = ratio (3.02). Hindwing length (1.02 cm): width (0.31 cm) = ratio (3.29).

ABDOMEN. — Green with median yellow band. Tergite 9 + ectoproct apodeme branched below
callus cerci; acumen of tignum, long slightly swollen apically; gonosaccus with numerous gonosetae;
arcessus apex quite curved, lacking dorsal striation; gonarcus with narrow lateral arms; gonosaccus
with numerous long gonosetae with crooked apices and ventral large gonocristae (Fig. 25C, D, E).

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Jaboticabal, 8 June 1995, Ferreira, R. J.
(50 paratypes)(guava), May 1995, Ferreira, R. J. (30% paratypes)(guava)(SDF/CAS).

Genus Chrysopodes Navas, 1913c

This is among the most diverse genera of New World Chrysopini. As with most genera in this
tribe, the genus is usually defined by elements of the male genitalia, notably the lack of a tignum and
gonapsis, presence of gonarcus with arcessus reduced to a curved, triangular, apically tapered sclerite.
The ventral branch of the dorsal apodeme of the ectoproct is often acutely tapered apico-medially.

This is a New World genus distributed from south Florida, U.S.A. and Baja California, Mexico
south to Argentina. The genus is usually divided into two subgenera (Chrysopodes and Neosuarius
Adams and Penny, 1987) based principally on the shape of the adult mandibles. Members of
Chrysopodes (Chrysopodes) have mandibles which are long and scythe-like, while members of
Chrysopodes (Neosuarius) have mandibles which are considerably stouter than the nominate subge-
nus (Adams and Penny 1987). In most adult specimens, the mandibles are tucked under the labrum
and difficult to see. The easiest way to view the mandibles is to separate the head from the body and
macerate it in 10% KOH. Because this disarticulates the body and makes it more difficult to view
other markings on the head, we have tried to avoid using mandibular characters in the species key.
Most of the species of C. (Neosuarius) found west of the Andes in South America are dark in color, but
species of C. (Neosuarius) found east of the Andes and in Central America are primarily green in
color. Species of C. (Chrysopodes) generally have proportionally broader wings, a broader costal
area, and longer forewing setae.

270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

Brooks and Barnard (1990) listed 21 species in C. (Chrysopodes) and 11 species in C.
(Neosuarius); Penny (1998) added two more species of C. (Chrysopodes), and Penny (2001) one addi-
tional species of C. (Neosuarius) from Costa Rica.

KEY TO THE SPECIES OF CHRYSOPODES OF BRAZILIAN AGRO-ECOSYSTEMS

1.. Meso-.and metanota with dark spots (Figs 29A) 3 <2.) <) s-cniS¢ Gdns) sexes oops Geel ee 2
1) Meso= and metanota without dark spotsi(Eigs 28A))= = gee.) es poise i ee) Cenc ee ee 4
2. One pair circular spots (Fig. 29A) or completely pale (Fig. 283A) on mesonotum; frons mostly pale, at most with
transverse dark stripe (Fig. 26€). 2. sco 5 erat Fe sopin a oes Fen ome epee «ee Sry en 3
2'. Irregular spots forming partial stripe on mesonotum (Fig. 30A); clypeus, gena, part of frons and lateral margin of
WEREXTTEG(EIPAS0G)) a oni cee oie Sin Soe cerca hag sy Segre Gen el cece eee ee C. (C.) copia

3. Antenna with flagellum black; dorso-medial stripe of scape incomplete, dark; two pair of lateral spots on pronotum;
mesopraescutum completely pale (Fig. 29A); forewing without crossvein shading (Fig. 29B).. . . . C (C.) adynatos
3'. Antenna with flagellum pale; dorso-medial stripe of scape reaching both proximal and apical margins; single pair
of elongate dark spots on pronotum, extending onto anterior margin of mesopraescutum (Fig. 34A); forewing

crossveins and gradates extensively shaded (Fig. 34B) ....:.....:.-....0.-5.0- C. (C.) nigropicta
4) sErons with semicircularred stripes below antennal|pits((E1g,26©)/e. sn een eee 8
4'=\Brons without stripes (Fig. 32A)) 22. 20... 80 a ee Re or 5
5. Gena with two longitudinal dark stripes; clypeus and labrum laterally dark (Fig.27D)................. 6
5/4 Gena completely redelypeus/and) labrum\pale((Bigs 36@))= = =) ee) Ul
6. Apical forewing veins of male swollen (Fig. 27E); lateral arms of gonarcus longer than width between arms

(Bige 27) ee eee Se ee ee ee te Pe hey Ce en Geet eee C. (C.) polygonica
6’. Apical forewing veins of male not swollen (Fig. 26B); lateral arms of gonarcus shorter than width between arms

(Big h32E) etre Cee ed eM ROR: SRE IL ee ee ae C. (C.) delicata
7. Scape with dark lateral spot (Fig. 36A); forewing with five inner and seven outer gradate veins (Fig. 36B);

spermatheca short, with short, uncoiled spermathecal duct (Fig. 36E).................. C. (N.) karinae
7'. Scape without dark lateral spot (Fig. 26A); forewing with three inner and six outer gradate veins (Fig. 31A);

spermatheca long and multiply-coiled, as is spermathecal duct (Fig.31F) ............... C. (C.) crocinus
8. Vertex and frons with dark marks between antennal bases (Fig.33C).................. C. (C.) elongata
8'. Vertex and frons without dark marks between antennal bases (Fig. 26A).....................-.. 9
9s Maletectoproctiwithitwo)ventrall projections|(E1g535D) mee eee) as ee cr ee ce 10

9'. Male ectoproct with single ventral projection, which is an extension of the dorsal apodeme (Fig. 261) C. (C.) lineafrons

10. Lateral arm of gonarcus longer than high (Fig. 271); small ventral peg on sternite 8 + 9 (Fig. 261) . . . C. (C.) spinella
10’. Lateral arm of gonarcus much higher than long (Fig. 35H); sternite 8 + 9 without ventral peg (Fig. 35D) C. (C.) divisa

Chrysopodes (Chrysopodes) lineafrons Adams and Penny, 1987

DIAGNOSIS. — Of the species with reddish bands below the antennae and dark palpi, C.
lineafrons males can be recognized by the strongly arched apex to the ninth sternite, the single ventral
projection from the ectoproct, and the triangular shape of the lateral arms of the gonarcus. Females can
be recognized by the heavily-sclerotized knob at the apex of sternite 7.

HEAD. — Vertex glossy green with two red marks lateral to central raised area (Fig. 26A). Frons
green with narrow red crescentic band below each antenna (Fig. 26C). Gena red, palest on central part.
Distal maxillary palpimere black; fourth palpimere dorsally black. Distal labial palpimere black (Fig.
26C). Scape and pedicel greenish yellow, unmarked; flagellum pale.

THORAX. — Green with yellow stripe along mid-line. Pronotum with lateral suffused pale brown
stripe (Fig.26A). Meso- and metanota unmarked. Wing: Forewing venation mostly green with some
transverse veins black, such as costal crossveins, R-Rs crossveins, base of Rs-Psm, inner and outer
gradates and apices of anal veins (Fig. 26B). Length (1.4 cm): width (0.77 cm) = ratio 3.0. Hindwing

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 27)

acute, venation green, except costal crossveins dark (Fig. 26B). Length (1.24 cm): width (0.39 cm) =
ratio 3.18.

ABDOMEN. — Green with yellow stripe on dorsal mid-line. Male: Dorsal apodeme of tergite 9 +
ectoproct heavily sclerotized with apical tooth projecting ventro-medially (Fig. 261). Apex of sternite
9 with conical-based setae (Fig.26E), ventrally, deeply concave on posterior half (Fig. 261). Gonarcus
narrow, strongly arched with long triangular lateral arms. Arcessus short, thin, evenly tapered to
pointed apex, which has heavily-sclerotized basal apodemes (Fig. 26F, G). Gonosaccus simple sac.
Female: Posterior margin of sternite 7 with heavily-sclerotized knob. Spermathecal duct short (Fig.
26H).

MATERIAL EXAMINED. — Brazil: Amazonas, Manaus, 17 February 1977, N.D. Penny,
holotype male (deposited at INPA, Manaus), S40 Paulo: Jaboticabal, 14 January 1999, Freitas, S.
(202 2)(guava); 26 April 1999, Takahashi, K. (1)(orange)(CAS); Sao Jose do Rio Preto, 20 April
1997, Freitas, S. (10); Taquaritinga, 5 March 1994, Xavier, A. L. (1 2)(CAS).

Chrysopodes (Chrysopodes) polygonica Adams and Penny, 1987

DIAGNOSIS. — The male can be recognized by the swelling of the forewing apical gradates. Al-
though males of other species of Chrysopodes have swollen veins, this usually involves the radial sec-
tor and associated veins more basal on the forewing. Only males of C. polygonica have an unswollen
radial sector and swollen gradates and apical forks. The female also has a unique characteristic—a
field of small setae on the knob at the apex of the seventh sternite.

HEAD. — Yellow with red and black markings. Gena with two black stripes continued onto lat-
eral border of clypeus (Fig. 27C). Frons with small black spot close to gena. Vertex with postocular
red spots at the posterior margin (Fig. 27A). Scape, pedicel and flagellum pale, unmarked. Maxillary
and labial palpi black (Fig. 27C).

THORAX. — Green with yellow stripe on mid-line. Pronotum with pale brown stripe laterally
(Fig. 27A). Meso- and metanotum without markings. Wings: Forewing venation green, except some
costal crossveins, R-Rs crossveins, first five Rs-Psm crossveins, gradates and marginal forks dark. All
transverse veins show at least a little shading. Apical gradate crossvein and apical branches of Rs
swollen in male wings; female gradates normal. Forewing length (1.40 cm): width (0.47 cm) = ratio
(2.98). Hindwing costal crossveins black; gradates black with brown shading (Fig. 27B). Length
(0.86 cm): width (0.27) = ratio (3.18).

ABDOMEN. — Green with yellow stripe on mid-line. Tergites unmarked. Male: Sternite 8 + 9
with knob at mid-length and apical setae on conical bases (Fig. 27G). Ventro-caudal margin of
ectoproct prolonged into an acute point. Large subanal plate densely setose. Gonarcus elongate, frag-
ile, slender. Arcessus elongate, digitiform, apically decurved. Gonosaccus dorsally with few long
gonosetae, and extended ventrally as lobe with numerous small setae (Fig. 27H, I). Female: Sternite 7
with terminal knob densely covered by small setae (Fig. 27F). Spermatheca bulbously inflated.
Spermathecal duct short (Fig. 27J). Subgenitale with small rounded posterior lobes and no anterior
prolongation (Fig. 27K).

MATERIAL EXAMINED. — Brazil: Amazonas, Manaus, Parque das Laranjeiras, 22 January
1981, J. R. Arias, holotype male (deposited at INPA, Manaus), S40 Paulo: Jaboticabal, 10 March
1999, Freitas, S. (1)(orange); 18 March 1998, Shinirara, A. (1 2); Mato Grosso: Itiquira, 14 Decem-
ber 1996, Scomparin, C. H. J. (1 2)(rubber); 11 January 1997, Freitas, S. (301 ?)(rubber); 13 January
1997, Freitas, S. (40°1 2)(rubber)(SDF/CAS); 20 January 1997, Freitas, S. (1 ?)(CAS); 23 February
1998, Freitas, S. (203 2)(rubber).

272 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

Chrysopodes (Chrysopodes) spinella Adams and Penny, 1987

DIAGNOSIS. — The double ventral projections of the ectoproct appear to associate this species
with C. collaris and C. divisa in Chrysopodes (Neosuarius) (Adams and Penny, 1987). However, C.
collaris has a much broader body and neither of these species have the unique knob at mid-length on
the ninth sternite. Chrysopodes (Chrysopodes) lineafrons has both subantennal red bands and knob at
mid-length of the ninth sternite, but has only a single ventral projection of the ectoproct. Females of
these four species can be separated only with great difficulty.

HEAD. — Yellow with red markings. Vertex glossy with red postocular spot from hind margin
forward only to mid-length (Fig. 28A). Frons yellow with interrupted red band extending from eye
margin medially below antennal sockets. Gena red. Maxillary distal palpimere black, fourth
palpimere dorsally black and third darkened. Labial distal and basal palpimeres black. Scape and
pedicel yellow, unmarked; flagellum stramineous (Fig. 28A, D).

THORAX. — Green with yellow stripe on mid-line. Pronotum longer than wide, with stripe
dorsolaterally. Meso- and metanotum unmarked (Fig. 28A). Wings: Wing venation mostly green, in-
cluding pterostigma. Forewing costal crossveins, basal R-Rs crossveins anterior and posterior inter-
sections, origin of Rs, gradates, marginal forks and anterior ends of Psm-Psc all dark (Fig. 28B);
forewing length (1.21 cm): width (0.41 cm) = ratio (2.95). Hindwing with costal crossveins and
gradates dark (Fig. 28B). Length (1.06 cm): width (0.34 cm) = ratio (3.12).

ABDOMEN. — Green. Male: Dorsal apodeme of ectoproct with two ventral projections. Ninth
sternite with small knob at mid-length. Gonarcus strongly arched horizontally, but with little vertical
arch. Lateral arms of gonarcus elongate and apically rounded. Arcessus short, triangular (Adams and
Penny 1987). Female: Spermatheca elongate with vela in form of crooked coil. Bursal duct elongate
and multiply coiled (Fig. 28D).

MATERIAL EXAMINED. — Brazil: Para, Sao Geraldo, 30 November to 8 December 1982, J. R.
Arais, holotype male (deposited at INPA, Manaus), Sao Paulo: Jaboticabal, February 1992,
Scomparin, C. H. J. (1 2)(orange).

Chrysopodes (Chrysopodes) adynatos de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 12-VII-96 Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — Only two other species of Chrysopodes in the region have dark spots on the
mesonotum. However, the spots on the other species are more diffuse and not as well delimited. The
thin central arch of the gonarcus is also distinctive, as are the dark antennae.

The name “adynatos” is taken from the Greek meaning weak, or without strength, and refers to
the thin medial arch of the gonarcus in this species.

HEAD. — Yellow. Gena, frons and vertex without marks. Scape with mid-dorsal, tapered black
spot; pedicel and flagellum black.

THORAX. — Pronotum pale yellow, with two pairs of black marks, each pair appears as short lat-
eral stripe (Fig. 29A). Mesonotum with pair of rounded black spots on scutum (Fig. 29A). Metanotum
unmarked. Wings: Forewing venation green, except costal crossveins, R-Rs crossveins, first Rs-Psm
veins; Psm-Psc crossveins and first two unforked marginal veins dark (Fig. 29B). Length (1.3 cm):
width (0.45 cm) = ratio (3.25). Hindwing venation green. Length (1.18 cm): width (0.34 cm) = ratio
(3.47).

ABDOMEN. — Yellow, unmarked. Male: Genitalia weakly sclerotized. Gonarcus thin with lat-
eral projections similar to gonocornua but originate from lateral arms of gonarcus. Arcessus longer
than wide, with broad medial decurved hook. Gonosaccus with long setae (Fig. 29C, D).

MATERIAL EXAMINED. — Known only from the holotype.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 273

Chrysopodes (Chrysopodes) copia de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP; FCAV Macadamia; 12/11/94; Freitas, S. SP70.”

DIAGNOSIS. — The most distinctive characteristic of this species is the almost completely red
coloration of the frons, as well as the extensive red coloration on meso- and metanota. The red color-
ation of the frons would appear to be an exaggerated extension of the transverse red bands found in
several other species. However, these other species have dark palpi, not pale palpi as in this species.

This species appears most closely associated with C. (Chrysopodes) indentata Adams and Penny,
1987, because of the pale palpi and especially the short, broad gonarcus and arcessus. However, they
can be separated because C. (Chrysopodes) copia has the almost completely red frons and clypeus, the
ventral bump at mid-length of the male ninth sternite is not as developed, and the gonarcus is more lat-
erally arched.

The name “copia” comes from the Latin for abundance or plenty and refers to the similarity to C.
indentata.

HEAD. — Golden yellow with red markings. Clypeus, frons and gena completely red, except for
restricted area below antennae sockets and between anterior tentorial pits (Fig. 30C). A red stripe runs
from gena to posterior eye margin laterad of antennal sockets (Fig. 30A). Vertex yellow, except later-
ally close to eye margin red. Scape, pedicel and flagellum pale without marks. Maxillary and labial
palpi pale, unmarked.

THORAX. — Slightly greenish yellow. Pronotum with diffuse red lateral stripes continuing onto
meso- and metanotum (Fig. 30A). Wings: Forewing green with some crossveins totally or partially
dark, such as posterior junctions of costal crossveins, R-Rs endings, Rs-Psm endings, inner and outer
gradates, Psm-Psc crossveins, apex of marginal forked and unforked crossveins; and apical Psm-Psc
crossveins shaded (Fig. 30B). Length (1.27 cm): width (0.43 cm) = ratio (2.95). Hindwing venation
mostly green, except costal crossveins at subcostal juncture and outer gradates dark. Length
(1.18 cm): width (0.36 cm) = ratio (3.28).

ABDOMEN. — Slightly greenish yellow, unmarked. Male: Ventral branch of dorsal apodeme of
tergite 9 + ectoproct quite large (Fig. 30D). Sternite 8 + 9 with knob on ventro-medial area (Fig. 30D).
Gonarcus evenly curved horizontally, barely curved vertically; lateral arms with long dorsal plate,
slightly upturned apically (Fig. 30F, G). Arcessus short, broader than long, with a field of quite short
setae distally on medial apex. Gonosaccus bearing scattered long setae dorsally and field of many
small setae ventrally. Female: Spermatheca vela twice coiled, then extended as a convoluted coil.
Bursal duct short, twisted (Fig. 30H); subgenitale bilobed without medial projection (Fig. 30E).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 22 January 1994, Freitas, S. (10
paratype)(on macadamia)(CAS); 22 February 1994, Freitas, S. (3o°l 2 paratypes)(corn); 22 February
1995, Freitas, S. (1 2 paratype)(guava)(CAS); 22 March 1995, Freitas S. (2 paratypes)(corn); 10 De-
cember 1996, Freitas, S. (1c paratype)(corn); Ribeirao Preto, October 1995, Pinotti, F. (2?
paratypes); Bebedouro, 14 October 1995, Fonseca, C. (1% paratype); Birigui, 11 July 1994,
Scomparin, C. H. J. (1o paratype)(corn).

Chrysopodes (Chrysopodes) crocinus de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo.Brazil, la-
beled “Birigui, SP, 21-II-96, Scomparin, C. H. J.” (corn).

DIAGNOSIS. — The most distinctive feature of this species is the lack of dark or red markings on
head and thorax. The male gonarcus is similar to most species in the genus. However, a few features
set this species apart. The dark antennal flagellum is found only in two other species of C.
(Chrysopodes)—C. (C.) adynatos and C. (C.) victoriae—both of which have dark or red markings on

274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

the thorax. Several South American species of C. (Neosuarius) have dark antennae, but again there are
extensive dark markings on both the head and thorax, as well as dark maxillary palpi.

There appears to be no ventro-medial point to the male ectoproct of C. (C.) crocinus, unlike most
species in this genus. (This character state could develop late in the maturation of the emerged adult).
Most distinctive is the basal point on the male sternite 8 + 9. Less easily discernible are the hirsute tip
of the arcessus and the two small patches of gonosetae on the gonosaccus.

The name “crocinus” comes from the Latin for saffronlike, referring to the coloration of the
antennal scape and pedicel.

HEAD. — Yellow. Frons, clypeus and vertex pale yellow, unmarked. Gena slightly darkened.
Scape, pedicel and basal membrane yellow-orange, without marks; flagellum black. Vertex without
postocular marks. Labial and maxillary palpi pale.

THORAX. — Green witha pale stripe on mid-line. Pro-, meso- and metathorax unmarked. Wings:
Forewing venation green, except costal crossveins and R-Rs crossveins somewhat darkened.
Intramedian cell ovate, with swollen base. Base of Psc near intramedian cell swollen with many
microtrichiae (Fig. 31C). Pterostigma unmarked. Length (1.18 cm): width (0.41) = ratio (2.88).
Hindwing acute; venation green; stigma unmarked. Length (1.12cm): width (0.35 cm) = ratio
(3.20)(Fig. 31A).

ABDOMEN. — Green with yellow stripe on middle line. Tergites unmarked. Male: Dorsal
apodeme of ectoproct not apically forked. Sternite 8 + 9 with knob near anterior margin (Fig. 31B).
Arcessus short, straight, triangular with dorsal setae most abundant. Gonosaccus with two groups of
three setae, each with large conical base. Gonarcus wider than long, with thin, broadly lobate, parallel
arms (Fig. 31D, E). Female: Sternite 7 with apical knob-like projection (Fig. 31G). Spermatheca long,
coiled. Spermathecal gland and bursal accessory glands long, coiled (Fig. 31F). Subgenitale saccate
(Fig. 31H).

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Birigui, 2 February 1996, Scomparin,
C.H. J. (2012 paratypes).

Chrysopodes (Chrysopodes) delicata de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo. Brazil, la-
beled “Itiquira, MT, Seringueira, PEM-507, 31/12/96 Scomparin, C. H. J. Armadilha—melaco”
(rubber).

DIAGNOSIS. — The double dark genal stripes, lack of subantennal bands, and dark palpi link C.
delicata with C. polygonica. However, males of C. polygonica have much longer lateral arms of the
gonarcus and swollen gradate and apical endings of forewing veins.

The name “‘delicata” comes from the Latin for delicate, tender, or dainty, and refers to the small
size of the gonarcus.

HEAD. — Yellow. Frons and vertex pale yellow, unmarked. Gena with two large black stripes
continuing onto margin of clypeus (Fig. 32A, C). Maxillary palpimeres |—4 and half of fifth black.
Scape, pedicel and flagellum pale (Fig. 32A). Mandibles scythe-like (Fig. 32E).

THORAX. — Green with yellow stripe on mid-line. Pro-, meso-, and metanotum unmarked.
Wings: Forewing green, except most crossveins dark and with slight shading; gradates more shaded
than other crossveins. Pterostigma green, unmarked. Forewing length (1.10 cm): width (0.38 cm) =
ratio (2.89). Hindwing costal crossveins, gradates and origin of marginal forks dark (Fig. 32B).
Length (0.96 cm): width (0.29 cm) = ratio (3.31).

ABDOMEN. — Green with yellow stripe on mid-line. Tergites unmarked. Male: Posterior margin
of ectoproct with long setae on thick bases. Sternite 8 + 9 with large knob at midlength and narrowed
apex (Fig. 32D). Male genitalia lightly sclerotized. Lateral arms of gonarcus reduced to tiny plates.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 2I3

Arcessus more sclerotized than other genital structures; apex smooth, without setae, decurved.
Gonosaccus with field of long setae (Fig. 32F,G).
MATERIAL EXAMINED. — Known only from the holotype.

Chrysopodes (Chrysopodes) elongata de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil
“Luiz Antonio, SP, Faz. Guatapara, 29-IV-94, Freitas, S.” (eucalyptus).

DIAGNOSIS. — This species is a member of a group of species with interrupted transverse band
across the frons and dark palpi. Chrysopodes elongata can be separated from C. lineafrons and C.
spinella by the small dark spots between the antennal bases. Additionally, females can be separated by
the long, narrow sternite 7 with no knobs, and especially by the long, thin spermatheca.

The name “elongata” comes from the Latin for prolonged, and refers to the long and narrow
spermatheca.

HEAD. — Yellow, marked with red. Frons yellow with narrow, medially interrupted, transverse
red line below antenna sockets; transverse line continued along ocular margin to posterior margin of
vertex. Gena red. Tiny red spot between antenna on frons and anterior part of vertex. Scape, pedicel
and flagellum pale. Fourth maxillary palpimere and half of fifth palpimere dark. Labial palpimeres
dark (Fig. 33A, C).

THORAX. — Green with yellow stripe on mid-line. Cervical sclerite red. Pronotum with pale
brown stripe dorso-laterally (Fig. 33A). Meso- and metanotum unmarked. Wing: Venation green, ex-
cept costal crossveins, R-Rs crossveins, gradates, origin of Rs, Rs-Psm crossveins and marginal forks
dark (Fig. 33B). Forewing pterostigma unmarked. Length (1.10 cm): width (0.38 cm) = ratio (2.89).
Hindwing costal crossveins and gradates dark. Length (0.97 cm): width (0.30 cm) = ratio (3.23).

ABDOMEN. — Green with yellow stripe on mid-line. Tergites without markings. Female:
Sternite 7 narrowed at apex. Spermatheca slender; duct extends as thin, filamentous tube to bursa
copulatrix (Fig. 33D, F). Subgenitale with small, medial projection (Fig. 33G).

MATERIAL EXAMINED. — Known only from the holotype.

Chrysopodes (Chrysopodes) nigropicta de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo Brazil,
labeled “Bra-MT-Itiquira, P. E.Michelin; 18/VIII/98, Freitas, S. 783A” (rubber).

DIAGNOSIS. — The black marks on pro- and mesonotum are unusual, as is the heavy shading of
forewing crossveins. Of the species having dark markings on the mesonotum, C. adynatos has dark
antennae and well delimited dark mesonotal spots. Although C. indentata also has pale antennae, the
dark markings of pro- and mesonotum are not found in this species. However, the two species appear
related because they both have an exceptionally heavily-sclerotized spermathecal duct.

The name “nigropicta” comes from the Latin nigra for black or dark and pictus for painted or col-
ored, referring to the dark markings of pro- and mesonota.

HEAD. — Greenish yellow. Vertex green, unmarked. Frons, clypeus and gena yellow. Scape yel-
low with mid-dorsal black stripe; pedicel and flagellum pale. Maxillary and labial palpi pale (Fig.
34A).

THORAX. — Green; pronotum with lateral black stripes from middle to posterior margin.
Mesonotum with pair of dark spots laterally on prescutum and second pair of lateral circular spots on
middle of scutum. Metanotum unmarked (Fig. 34A). Wings: Forewing veins green, with costal
crossveins, R-Rs crossveins, first Rs-Psm crossvein, gradates, Psm-Psc crossveins after intramedian
cell, apex of marginal forks, anterior marginal unforked veins and endings of anal veins dark. Almost
all crossveins in posterior half of forewing heavily shaded. Length (1.41 cm): width (0.5 cm) = ratio

276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

(2.83). Hindwing venation mostly green, except middle part of Rs and gradates dark (Fig. 34B).
Length (1.24 cm): width (0.40 cm) = ratio (3.10).
ABDOMEN. — Greenish yellow. Female: Spermatheca U-shaped. Spermathecal gland with dis-
tal part exceptionally heavily sclerotized. Bursa sac-like, with two accessory glands (Fig. 34C).
MATERIAL EXAMINED. — Known only from the holotype.

Chrysopodes (Neosuarius) divisa (Walker, 1853)

DIAGNOSIS. — This is one of several species which has medially interrupted, red bands across
the frons below the antennae. The double ventral projection of the ectoproct and tiny setae at the tip of
the arcessus can be found in the closely related C. collaris (Schneider, 1851). However, C. collaris isa
larger, thicker-bodied species with pronotum much wider than long; the apex of the sternite 8 + 9 is
truncated and vertically thick, and the gonarcus + arcessus complex has an H-shaped supporting brace
not found in C. divisa (see Adams and Penny 1987).

HEAD. — Yellow with red markings. Vertex with red lateral postocular marks from mid-length
to posterior margin. Frons yellow with red band below antennal bases interrupted along mid-line.
Gena red. Maxillary distal palpimere black; fourth palpimere dorsally black; and third fuscous. Third
labial palpimere black. Scape and pedicel yellow, unmarked; flagellum pale yellow (Fig. 35A, C).

THORAX. — Green with yellow stripe on mid-line. Pronotum longer than wide, with weakly de-
fined brown stripe laterally. Meso- and metanotum unmarked (Fig. 35A). Wings: Venation green, ex-
cept costal crossveins, origin of Rs, R-Rs crossveins, gradates; Psm-Psc crossvein endings; and
bifurcations of forked marginal veins dark. Pterostigma green, unmarked. Length (1.14 cm): width
(0.39 cm) = ratio (2.92). Hindwing venation green except gradates and middle costal crossveins dark
(Fig. 35B). Length (0.88 cm): width (0.28 cm) = ratio (3.14).

ABDOMEN. — Green, with mid-line yellow. Tergites without markings. Male: Tergite 9 +
ectoproct triangularly angled at posterio-ventral margin, with basal lobe and apico-medially project-
ing hook (Fig. 35D). Sternite 8 + 9 deeply constricted posteriorly, without knob, with lateral setae
with thick bases. Gonarcus mildly arched, with oblong, curved lateral arms. Arcessus evenly tapered,
triangular, with apex decurved and bearing tiny setae (Fig. 35G, H). Female: Dorsally folded bursa
with pair of ovate accessory glands. Long bursal duct extended into long and narrow spermatheca
(Fig. 35E, F). Subgenitale broader than long, with small medial projection.

MATERIAL EXAMINED. — Brazil: Sao Paulo: Taquaritinga, 6 August 1993, Xavier, A. L. Q.
(1o)(orange); 8 January 1993, Xavier, A. L. Q. (1o%)(orange); 5 March 1994, Xavier, A. L. Q.
(40°1 $)(orange)(SDF/CAS); 19 November 1992, Freitas, S. (22 )(orange); 10 January 1994, Xavier,
A. L. Q. (1 ?)(orange); 5 March 1994, Xavier, A. L. Q. (142)(orange); Monte Azul Paulista, Novem-
ber 1994, Silva, J. L.(1@)(orange); Barretos, 1 1 February 1991, Freitas, S. (1 o)(orange); Jaboticabal,
20 August 1995, Freitas, S. (12); Luiz Antonio, 21 December 1994, Freitas, S. (1 2)(eucalyptus);
Elisario, July 1994, Silva, J. L. (1 2)(orange); Minas Gerais: Peti, October 1988, coll. not named
(itch);

Chrysopodes (Neosuarius) karinae de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Bra-SP-Jaboticabal, FCAV, 24/IV/99, Takahashi, K. M., 822” (guava).

DIAGNOSIS. — This species is unlike other green species of Chrysopodes (Neosuarius) in having
antennae with dark flagella, scapes with lateral mark, and labial and maxillary palpi pale. Itcan also be
separated from C. collaris by having a pronotum as long as wide (Adams and Penny 1987). However,
the most unique characteristic of this species is the subanal plate with numerous small setae and lateral
pockets, which is found in no other species. The spermatheca is also thinner and more elongate than

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 20h

most species. The spermathecae shape, long and crooked, the setae around the genitalia atria and the
lateral pocket are not found in any other chrysopid species.

The name was given in honor of Agricultural Engineer Karina Manami Takahashi, enthusiastic
entomologist who has studied and collected many chrysopids.

HEAD. — Golden yellow with red markings. Frons and clypeus unmarked. Gena red-tinged. Ver-
tex with red spots on posterior margin near eyes. Scape yellow with dark spot at dorso-lateral apex;
pedicel and flagellum dark (Fig. 36A, C). Maxillary and labial palpi pale.

THORAX. — Yellow. Pronotum with lateral pale brown stripe (Fig. 36A). Meso- and metanotum
unmarked. Wings: Forewing venation green, except margin endings costal crossveins, R-Rs
crossveins and gradates dark. Length (1.45 cm): width (0.5 cm) = ratio (2.9)(Fig. 36B). Hindwing ve-
nation green. Length (1.33 cm): width (0.43 cm) = ratio (3.09).

ABDOMEN. — Yellow, marked red laterally on first tergite. Female: Tergite 8 ven-
trally-prolonged laterally. Tergite 9 + ectoproct dorsally raised. Sternite seven with ventral knob api-
cally (Fig. 36D). Proximal to genital atria a dorsal field of setae, a medial plate with densely clumped
setae and two lateral pockets (Fig. 36F). Spermatheca elongate, twisted but not coiled, with long duct
connected to bursa copulatrix. Bursa a membranous sac with pair of ovate accessory glands (Fig.
36E). Subgenitale simple, apically bilobed.

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo, Jaboticabal, 24 April 1999, Takahashi,
K. M. (1% paratype)(guava).

Genus Plesiochrysa Adams, 1982b

Plesiochrysa adults are among the largest of South American Chrysopini, with forewing length
of about 15 to 18 mm. They can usually be found in open fields, and at times can be quite common.
Their male genital elements indicate a close relationship with the Holarctic genus Chrysopa. Brooks
and Barnard (1990) listed 23 species in this genus, distributed in the Neotropics, Australia, Malay Ar-
chipelago, southern India to the Seychelles. Three species have been found during this study, includ-
ing one new species. Unfortunately, male genital elements of all Brazilian species are virtually
identical, but the species can be identified using the following key.

KEY TO THE SPECIES OF PLESIOCHRYSA FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

lmeAntennacidarka(EipasQA)i see 6 coal ho rie cose el ie eh ate Gl gh Gna eae Plage voll RM eh Seek eek P. alytos
IMATItcOnac Pale (IG SSA er. cue bins a eked sn cat is. bese ae pews: wade a Aackee kk Voll bbs wah heap oe omens 2
2. Pronotum longer than wide; lateral stripes continuous or only slightly interrupted (Fig. 38A) ........ P. elongata
2'. Pronotum as wide as long; lateral spots widely separated (Fig. 37A)...................4. P. brasiliensis

Plesiochrysa brasiliensis (Schneider, 1851)

DIAGNOSIS. — Plesiochrysa brasiliensis can be separated from the other two Brazilian species
(P. elongata and P. alytos) by the dark antennal flagellum of P. alytos and longer pronotum with more
continuous lateral red markings of P. elongata.

HEAD. — Yellow with red marks. Vertex yellow with small red spots on posterior part near eye
margin (Fig. 37A). Frons unmarked. Gena with narrow red stripe close to frons continued onto lateral
margin of clypeus (Fig. 37C). Maxillary palpi with dorsal black stripe. Labial palpi pale, unmarked.
Scape and pedicel pale, unmarked; flagellum pale yellow.

THORAX. — Green with pale yellow stripe on mid-line. Pronotum marked with pair of
well-delimited red spots laterally (Fig. 37A). Meso- and metanota unmarked. Wings: Forewing vena-
tion green, except posterior junctions of costal crossveins 3—7 and 8—13, origin of Rs, R-Rs crossveins

278 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1-4, gradates, Psm-Psc crossveins, marginal unforked veins near anals and apices of anal veins dark
(Fig. 37B). Length (1.5 cm): width (0.5 cm) = ratio (2.98). Hindwing venation green, except outer
gradates dark (Fig. 37B). Length (1.30 cm): width (0.43 cm) = ratio (3.02).

ABDOMEN. — Green with yellow stripe on mid-line. Male:Tergite 9 + ectoprocts widely sepa-
rated with medial indentation (Fig. 37D). Sternite 8+ 9 elongate, narrowed at apex (Fig. 37E).
Tignum narrow, transverse band. Gonarcus wide and thick with small, medial gonocornua.
Entoprocessus not fused to gonarcus, Y-shaped. Arcessus absent. Pseudopenis elongate, tapering,
apically decurved. Gonosaccus large with numerous long setae on tubercules (smaller near
pseudopenis) (Fig. 37G, H). Female genitalia: Spermatheca short. Bursa sac-like (Fig. 37F).
Subgenitale with lateral lobes and small median projection.

MATERIAL EXAMINED. — BRAZIL: Sao Paulo: Jaboticabal, November 1991, Hungaro, C. M.;
24 October 1992, (12); Tanigawa, F. (12); October 1992, Mendes, M. B. (12); October 1992,
Tannuri, L. (1<); 5 November 1992, Scomparin, C. H. J. (1 2)(orange); 22 November 1994, Freitas,
(1o)(guava); 30 June 1995, Freitas, S. (10°); 26 January 1995, Freitas, S. (10°); 22 February 1995,
Freitas, S. (1o°l 2 (orange); 24 February 1996, Rossi, R. M. (1 ?); 28 August 1996, Silva, L. C. (1¢);
20 October 1996, Freitas, S. (1o)(guava)(CAS); 21 October 1996, Fernandes, M. (12); 20 October
1996, Freitas, S. (2o)(orange); 9 December 1996, Freitas, S. (1o°)(orange)(CAS); 17 June 1998,
Freitas S. (10°); Birigui, 26 January 1996, Scomparin, C. H. J. (171 2)(corn); Atibaia, 15 November
1994, Onara, L. (1 2); Eng. Coelho, 10 August 1996, Poletti, M. (10°); Matao, 15 October 1996, Yiagi,
R. (1c); Olimpia, 2 August 1996, Tolfo, A. L. T. (12).

Plesiochrysa elongata (Navas, 1913b)

DIAGNOSIS. — This species can be separated from P. a/ytos by its pale antennae and pale gradate
crossveins, and from P. brasiliensis by its longer pronotum with more complete stripes.

HEAD. — Yellow with red markings. Vertex yellow with tiny red spots beside eye at posterior
margin. Frons unmarked. Gena with red stripe on half closest to frons, not extended onto clypeus (Fig.
38A, C). Maxillary palpi with dorsal black stripe. Labial palpi unmarked. Scape, pedicel and
flagellum pale yellow, unmarked.

THORAX. — Green with pale yellow stripe on mid-line. Pronotum marked with red stripe nar-
rowed or interrupted at midlength. Meso- and metanota unmarked (Fig. 38A). Wings: Venation and
size similar to P. brasiliensis.

ABDOMEN. — Male and female as P. brasiliensis.

MATERIAL EXAMINED. — BRAZIL: Sao Paulo: Jaboticabal, October 1993, Furuhashi, S. (1<);
10 October 1996, Ferreira, A. (1); 15 November 1996, Freitas, S. (1 2)(corn)(CAS); October 1993,
Fernandes, F. (1 2); Minas Gerais: Vicosa, 28 December 1988, Abrantes, C. V. M. (11 2); 27 Janu-
ary 1989, Abrantes, C. V. M. (2o&°)(SDF/CAS); 28 February 1988, Abrantes, C. V. M. (10); Mato
Grosso: Itiquira, 10 September 1996, Scomparin, C. H. J. (1 2)(rubber).

Plesiochrysa alytos de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, PEM, 7-IX-99, Freitas, S.” (guava).

DIAGNOSIS. — This is the only species of Plesiochrysa in South America which has a continuous
pronotal band, showing no signs of restriction or broken lines at mid-length. In the other species, occa-
sional specimens will appear to have stripes rather than spots, sometimes because of pigment “bleed-
ing,” but there is virtually always an indication of restriction at mid-length. This is also the only
species with dark antennae.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 259

This name “alytos” comes from the Greek, meaning continuous or unbroken, and refers to the
continuous, unbroken, lateral stripes on the pronotum.

HEAD. — Yellow with red markings. Vertex yellow with small red spots beside eye at posterior
margin (Fig. 39A). Frons and clypeus unmarked. Gena red-tinged. Maxillary palpi with dorsal black
stripe. Labial palpi unmarked (Fig. 39C). Scape and pedicel unmarked, flagellum black basally and
fuscous apically (Fig. 39A).

THORAX. — Greenish yellow. Pronotum longer than wide, glabrous, with pale red lateral stripe.
Wings: Venation green, except darkened gradates in fore- and hindwings (Fig. 39B). Forewing length
(1.56 cm): width (0.49 cm) = ratio (3.18). Hindwing length (1.40 cm): width (0.43) = ratio (3.26).

ABDOMEN. — Greenish yellow. Male and female genitalia, as in P. brasiliensis.

OTHER MATERIAL EXAMINED. — BRAZIL: S4o Paulo: Jaboticabal, November 1996, Manzini,
J. P. (12 paratype); 1 November 1990, Camargo, R. (1o paratype); 13 November 1994, Lopez, C. C.
(101 2 paratypes)(SDF/CAS); 12 September 1995, Gomes, R. G. (1% paratype); 20 November 1995,
Ferreira, R. J. (lo paratype); 12 September 1996, Britto, L. F. (12 paratype)(CAS); 4 December
1996, Freitas S. (12 paratype); Birigui, 26 January 1996, Scomparin, C. H. J. (1? paratype)(corn);
Pirangi, 15 December 1996, Fernandes, E. (1 @ paratype); Franca, October 1990, Cunha, L. S. A. (1¢
paratype); Mato Grosso: Itiquira, 4 November, 1996, Scomparin, C. H. J. (1o paratype)(rubber); 7
September 1996, Freitas, S. (1% paratype)(rubber).

TRIBE LEUCOCHRYSINI ADAMS, 1978

Adult members of the tribe Leucochrysini are distinctive. They have a dark spot along the ante-
rior margin of the forewing at the base of the pterostigma, a rather low angle of juncture between the
pseudomedia and the outer gradate series of crossveins, and antennae which are longer than the wings.
Males have a broad gonarcus which is only weakly arched and lacks entoprocesses. Female genitalia
often include an elongate subgenitale, which is likely the precursor for a praegenitale (Brooks and
Barnard 1990).

This tribe is only found in the New World, ranging from as far north as lowa and Massachusetts in
North America (Penny et al. 1997) south to Argentina (Penny 1978). Members of Leucochrysini are
found primarily in tropical forests where they are the dominant tribe in this habitat. Leucochrysa, with
its two subgenera, is the most speciose genus of the family. Leucochrysini consists of seven genera,
and is most diverse in South America. All genera are known from Brazil, except Neula, which is
known only from its type specimen collected in Colombia. All known larvae are trash-bearers, known
locally as “‘bicho de lixo.”’ Probably because of their preference for forest habitat, only Leucochrysa
has been collected in Brazilian agro-ecosystems.

Genus Leucochrysa McLachlan, 1868

Much of what has been said about Leucochrysini applies equally well to Leucochrysa. It is the
largest genus in the family with nearly 170 described species and many still undescribed. Like the
tribe, Leucochrysa is distributed from northern U.S.A. to Argentina, although the greatest diversity
and numbers occur in the tropical forests. Within the tribe, Leucochrysa is currently delimited by the
lack of diverse derived character states of the other genera. A phylogenetic analysis of the tribe will
probably split Leucochrysa into several lineages. As currently constituted, Leucochrysa has two sub-
genera. Adults can be distinguished using the following key.

280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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KEY TO THE SUBGENERA OF LEUCOCHRYSA

1. Forewing intramedian cell quadrangular, Rs straight (Higs40A)i 9 SRR ee ee L. (Leucochrysa)
1”, Forewing intramedian cell| triangular, Rs sinuous (Fig. 47B)), 22 2-9-4) 2 2 2 L. (Nodita)

Subgenus Leucochrysa (Leucochrysa) McLachlan, 1868

This subgenus is usually characterized by having forewings with a quadrate intramedian cell,
rather than an ovate or triangular one, and straight radial sector rather than sinuous. However, as is fre-
quently pointed out (Banks 1945; Adams 1977; Brooks and Barnard 1990), a small percentage of indi-
viduals of either subgenus will have the shape of the intramedian cell of the other subgenus.
Individuals of Leucochrysa (Leucochrysa) are large, with a forewing length of more than 20 mm and
at times more than 30 mm. The shape of the intramedian cell may in part be correlated with large wing
size, although large species of Leucochrysa (Nodita) maintain a triangular shape. The shape of the
forewing radial sector is also probably an artifact of larger wings of Leucochrysa (Leucochrysa), as
larger species of Leucochrysa (Nodita) also appear to have a straighter Rs. Brooks and Barnard (1990)
recorded 41 species in Leucochrysa (Leucochrysa) distributed from northern U.S.A. to Argentina.
Seven species are recorded here, two were previously undescribed. Adults can be distinguished using
the following key.

KEY TO SPECIES OF LEUCOCHRYSA (LEUCOCHRYSA) FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

1. Two longitudinal dark stripes on meso- and metanota; pronotum with two red spots near posterior margin

(Big SUA Ar eS re a de ace a et ee, Ba ole Nr L. (L.) boxi
1’. Meso- and metanota pale or dark, but without longitudinal stripes; pronotum pale green without markings

(Fig: 40C).. 4a aon ee oo ee ee ee ee be ee Oe 6 Te eee 2
2) Headhthoraxsandiabdomen without markings. 9) sie) Sciences see) ee L. (L.) catarinae
2'. Head with thick red markings and suffusions (Fig. 40B, C); abdomen with large dark marks on Tergites 5 and 6

(E1gsAOD) Ree cence tee tt ett anelye aeoay Subse re yor et codes shel ce Ge Spel EP ave oe ae er 3
32) Darksspotiat base of forewing intramedia\vein\(Figs40A)i 2 23). 4 os = ens ee) cee 4
3/4 Noidark/spot at baseiof forewing intramedia vein\(Fig.44A)" = 32 22 2.6) «2 eee eee 5
4. Frons suffused with red (Fig. 43D); female spermatheca coiled about five times, with short spermathecal duct

(hips 43B)eiy secon cree lee, Stic feo eaters ee dee ee. ee Se eee L. (L.) varia
4’. Frons pale, without markings (Fig. 40B); female spermatheca short, S-shaped, with long spermathecal duct coiled

dozens:ohtimes (BigZ40E)i = foe cerpce cones tes yh ich nee eae ee L. (L.) ampla
5. Male gonocornua short, separated by more than five times their length (Fig.44D) .......... L. (L.) walkeriana
5'. Male gonocornua longer, separated by less than twice their length (Fig. 42D). ..................... 6

6. Lateral arms of male gonarcus oriented vertically; inner margins of gonocornua diverging (Fig. 42D). . L. (L.) pretiosa
6’. Lateral arms of male gonarcus oriented at 45° to vertical; inner margins of gonocornua parallel (Fig. 45C)
Sop PE Oey (CoC OCM EMO Note RY COMER ON Te ob CMe Co ad MCMC Ome McEwen HMMM MS oe Sc L. (L.) bruneola

Leucochrysa (Leucochrysa) ampla (Walker, 1853)

DIAGNOSIS. — This is a member of a group of species having heavy red markings and red pig-
mentation suffused over the frons and vertex, no markings on the pronotum, general (but variable)
dark suffusion of the meso- and metascuta, and dark marks on the Sth and 6th abdominal tergites.
Within this group, L. ampla can be separated from all others, except L. varia, by the dark spot at the
base of the intramedian cell. From L. varia it is separated by distinctly different female genitalia: L.
varia having a long, multiply coiled spermatheca and short, uncoiled spermathecal duct, while L.
ampla has an S-shaped spermatheca and long spermathecal duct coiled three to four dozen times.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 281

HEAD. — Pale yellow, broadly tinged reddish brown; vertex reddish along posterior ocular mar-
gin, lateral of raised medial area and base of antenna; frons dark brown between scape and eye and
through antennal sockets to ventral margin with gena; gena dark brown at frons and clypeus and other-
wise red; scape light red-tinged throughout; flagellum pale, basal segments fuscous; maxillary and la-
bial palpi pale (Fig. 40B).

THORAX. — Cervical sclerites red; pronotum pale yellow, unmarked; most of meso- and
metanota dark red and chocolate brown (Fig. 40C). Wings: Pterostigma marked with prominent dark
brown spot. Forewing bases of longitudinal veins dark brown; outer gradates with conspicuous brown
spots; inner gradates brownish; most costal cross veins brown at apex; base of Rs and ventral corner of
intramedian cell brown with bordering; marginal forks of veins brown at apex. Length (2.11 cm):
width (0.83cm) = ratio (2.54). Hindwing veins predominately pale, only costal crossveins brown (Fig.
40A). Length (1.87 cm): width (0.63 cm) = ratio (2.97).

ABDOMEN. — Yellowish green with a broad yellow stripe along dorso-medial line; tergites 5 and
6 dark brown (Fig. 40D). Female genitalia: Spermatheca long; spermathecal duct long, convoluted;
subgenitale bears extra plate with pit; long accessory glands (Fig. 40 E, F).

MATERIAL EXAMINED. — BRAZIL: Mato Grosso: Itiquira, 13 January 1997, Freitas, S.
(1 2)(rubber).

Leucochrysa (Leucochrysa) boxi Navas, 1930

DIAGNOSIS. — The characteristic of a medial dark stripe on the meso- and metanota is shared by
L. (L.) longicornis and L. (L.) boxi. In characters of the male genitalia they are also similar, including
the subapical constriction of sternite 8 + 9. However, the two species differ in that L. (L.) longicornis
has a tapering dark medial stripe on the pronotum, while L. (L.) boxi is pale medially and has two short,
dark, lateral marks on the pronotum. The inner gradate series of the forewing of L. (L.) boxi is also dis-
tinctive, extending quite far basally and parallels the pseudomedius for a long distance.

HEAD. — Yellow; frons, clypeus, vertex and gena pale; scape and pedicel dark laterally and
dorso-laterally; first 17 antennomeres (including pedicel) black on anterior surface; maxillary and la-
bial palpi pale.

THORAX. — Pronotum dark at anterio-lateral margin; meso- and metanota dark at mid-line (Fig.
41A). Wings: Venation mostly green; pterostigma dark; intramedian cell quadrangular; inner
gradates curved medially to level of origin of fifth radial cross veins; dark oblique band at wing apex.
Length (2.42 cm): width (0.84 cm) = ratio (2.88). Hindwing costal crossveins black, remainder of
wing green (Fig. 41B). Length (2.21 cm): width (0.71 cm) = ratio (3.11).

ABDOMEN. — Dorsal surface and tergites dark-tinged (Fig. 41C). Male genitalia: Gonarcus lat-
eral arms broad; gonarcus hardly curved and thick; gonocornua short, upturned with broad base;
arcessus short with lateral lobes, medial lobe of arcessus decurved; gonosaccus without gonosetae
(Fig. 41E, G). Dorsal apodeme of ectoproct forked posteriorly (near callus cerci)(Fig. 7D); sternites
with many microtholi; sternites 8 + 9 narrow at apex, bearing setae (Fig. 41F).

MATERIAL EXAMINED. — BRAZIL: Santa Catarina: Sao Joaquim, 20 August 99, Borges, R.
(1%)(apple).

Leucochrysa (Leucochrysa) pretiosa (Banks, 1910)

DIAGNOSIS. — This species forms part of the varia species group having extensive red suffusion
of the frons and vertex, completely pale pronotum, and heavy, dark, poorly delimited markings of the
meso- and metanota. It is one of three species having no dark spot at the base of the intramedian cell.
Amongst these three species male genital characters are the most reliable. Leucochrysa bruneola has
lateral arms of the gonarcus angled at about 45° and closely spaced gonocornua which are short and

282 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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parallel. Leucochrysa walkeriana has vertically oriented lateral arms of the gonarcus, but the short,
parallel gonocornua are widely separated. Leucochrysa pretiosa has lateral arms of the gonarcus ver-
tically oriented, and more massive, closely-spaced gonocornua which appear to diverge at their inner
margins.

HEAD. — Yellow, but broadly red-tinged; vertex wine red, raised medial area pale, between
antennal sockets brown-tinged; frons, clypeus and labrum wine red with some clear areas; gena red;
maxillary and labial palpi pale; scape and pedicel red; flagellum pale (Fig. 42A).

THORAX. — Cervical sclerites red; pale pronotum yellow unmarked; most of mesonotum brown;
metanotum brown except scutellum and borders of scutum (Fig. 42E); pterostigma faintly black
marked. Wings: Forewing black-tinged basally; longitudinal veins (costa, Rs, pseudomedia,
pseudocubitus, and anals) black at base; inner and outer gradates black; four apical Psm—Psc
crossveins black and outer gradates with brown shading; basal Rs crossveins black at intersections.
Hindwing venation mostly pale; costal crossveins black. Forewing length (2.03 cm): width (0.72 cm)
= ratio (2.82). Hindwing length (1.81 cm): width (0.60 cm) = ratio (3.02)(Fig. 42B).

ABDOMEN. — Green, dorso-medial line pale yellow, tergites 5, 6 marked with brown (Fig. 42J);
Male: Ventral fork of dorsal apodeme of ectoproct long, with acute apex (Fig. 42C). Gonarcus
slightly arched, lateral arms large, vertically oriented (Fig. 42G); gonocornua slender, decurved;
arcessus medial point decurved with thin lateral lobes, not seen in dorsal view; few gonosetae on
gonosaccus (Fig. 42D, G). Female: genitalia with convoluted spermatheca, vela large, gland short;
two long filamentous bursal glands present (Fig. 42F); subgenitale divided into two parts (Fig. 42H,
I); two sclerotized plates close to base of subgenitale with cerci and two irregular sclerotized struc-
tures can be seen (Fig. 3H, I).

MATERIAL EXAMINED. — Venezuela: San Esteban, | January, Anduze, syntype male (depos-
ited in Museum of Comparative Zoology, Harvard University), Brazil: Mato Grosso: Itiquira, 23
January 97, Freitas, S. (30¢22)(rubber); 5 April 97, Freitas, S. (1o)(rubber); 4 November 96,
Scomparin, C. H. J. (1@)(rubber).

Leucochrysa (Leucochrysa) varia (Schneider, 1851)

DIAGNOSIS. — Leucochrysa varia forms part of a species complex with character states men-
tioned above for other species. The dark spot at the base of the intramedian cell help distinguish this
species from all others in the complex, except L. ampla. The most distinctive characteristics separat-
ing these two species are the multiply-coiled spermatheca and short, uncoiled spermathecal duct of L.
varia.

HEAD. — Yellow with red markings. Vertex, frons, clypeus, genae, labrum, scape and pedicel
marked pale red. Maxillary and labial palpi unmarked (Fig. 43B, D).

THORAX. — Cervical sclerite red; pronotum yellow unmarked; meso- and metanotum
brown-tinged (Fig. 43B). Wings: Pterostigma faintly black marked; marginal endings of costal
crossveins black; apex of Rs dark; inner gradates pale, outer gradates black and shaded, first and sec-
ond ones more evident; third and apical psm-pscu crossveins black and shaded (Fig. 43A); length
(2.08—2.13 cm); width (0.72—0.83). Hindwing middle costal crossveins, black; apex of Rs faintly
dark; length (1.83—1.86 cm); width (0.54—0.56 cm) (Fig. 43A).

ABDOMEN. — Yellowish, 5th and 6th tergites marked with brown (Fig. 43C). Male: Dorsal
apodeme of ectoproct well-sclerotized and prolonged posteriorly into an acute apex; few microtholi
present; callus cerci with few trichobothria, whose setae are strong and short (Fig. 43E); Male genita-
lia with stout gonarcus and vertically elongate lateral arms; gonocornua short, not extended to apex of
arcessus; arcessus a heavily sclerotized sac from which apex of gonocornua extends; apical lobes lat-
erally rounded, medial hook decurved, heavily sclerotized; gonosaccus with a few gonosetae (Fig.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 283

43G, H). Female: Genitalia with spermatheca long and convoluted; gland short; bursa sac-like (Fig.
43E); subgenitale bilobed, with short ventral projection (Fig. 43F).

MATERIAL EXAMINED. — Brazil: unknown locality, holotype male (deposited in Museum of
Comparative Zoology, Harvard University), S40 Paulo: Jaboticabal, April 1993, Murtati, G. (1);
Luiz Antonio, 29 April 1994, Freitas, S. (1 2)(eucalyptus). Mato Grosso: Itiquira, 23 January 1997,
Freitas, S. (10)(rubber).

Leucochrysa (Leucochrysa) walkerina Navas, 1913

DIAGNOSIS. — This species is a member of the varia species complex. The lack of a dark spot at
the base of the intramedian cell groups it with two other species: L. bruneola and L. pretiosa. The most
distinguishing feature of L. walkeriana is the widely spaced gonocornua of the male genitalia. It also
has smaller gonocornua than L. pretiosa and more vertically aligned lateral arms of the gonarcus than
L. bruneolus.

HEAD. — Yellow, marked with red. Raised medial area of vertex with anterior converging
stripes extended to antennal bases; lateral area red-tinged. Scape and pedicel red-tinged; flagellum
pale, first five antennomeres with small black spot along anterior surface. Gena red. Frons with pair of
small red spots below antennae. Clypeus red-tinged medially (Fig. 44C).

THORAX. — Pronotum bright green, without marks. Meso- and metanota dark; mesonotum with
brown marks along anterior prescutum and lateral scutum (Fig. 44B). Wings: Venation pale, except
basal area, marginal junctions of costal crossveins, first radial crossvein, gradates, apical Psm-Pscu
crossveins dark. forewing length (2.15 cm): width (0.85 cm) = ratio (2.52). Hindwing costal
crossveins and middle part of Rs dark (Fig. 44A). Length (1.88 cm): width (0.67 cm) = ratio (2.81).
Pterostigma of fore- and hindwing with dark mark basally.

ABDOMEN. — Green, unmarked. Male: Gonarcus lateral arms slightly higher than long.
Gonocornua short, slightly decurved. Entoprocessus sharply decurved at base of gonarcus.
Arcessus broad with lateral lobes; medial hook apically decurved. Gonosaccus without gonosetae
(Fig. 44D, E).

MATERIAL EXAMINED. — Brazil: S40 Paulo: Campinas, 22 July 1991, Chinchilla, C. (1 0)(or-
ange).

Leucochrysa (Leucochrysa) bruneola de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 10-V-96, Seringueira, PEM 811, Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — This is amember ofa group of species having extensive red suffusion of the head,
completely pale pronotum, and extensive dark markings on the mesonotum. The lack ofa dark spot at
the base of the intramedian cell is characteristic of this and two other species: L. pretiosa and L.
walkeriana. The most distinctive feature of L. bruneola is the lateral arms of the gonarcus oriented at a
45° angle to vertical, as opposed to the more usual 90° angle in other species (Fig. 45C, E). Addi-
tionally, the gonocornua are more widely spaced in L. walkeriana, and gonocornua are larger and
more divergent along the inner margins in L. pretiosa (Fig. 42D).

The name “bruneola” comes from the Medieval Latin brunneus for brown and Greek olos for
mud or dirt and refers to the brown meso- and metanota.

HEAD. — Pale green. Gena red. Frons and clypeus lightly red-tinged. Vertex with red A-shaped
mark with legs expanded near eye margins; pair of small red spots along posterior margin near eye
(Fig. 45B). Scape and pedicel completely suffused with red; antennal fossa red.

THORAX. — Cervical sclerites red. Pronotum pale, unmarked. Meso- and metanota pale with
dark brown tinge; mesoprescutum and scutum with amorphous dark spots; second axillary sclerite

284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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dark (Fig. 45B). Wings: Forewing venation pale, costal crossvein apices, first four R-Rs crossveins,
gradates, and apical Psm-Psc crossveins dark. Length (2.1 cm): width (0.82 cm) = ratio (2.56).
Hindwing venation pale, except costal crossveins dark (Fig. 45A). Length (1.81 cm): width (0.63 cm)
= ratio (2.87). Pterostigma of both fore- and hindwing with dark basal spot.

ABDOMEN. — Yellow, unmarked. Male: Gonarcus thick, wide and short. Gonocornua conical,
apically decurved. Arcessus with heavily-sclerotized median, decurved, apical hook; lateral lobes not
projecting posteriorly. Gonosaccus with few large gonosetae (Fig. 45C, D, E).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Leucochrysa) catarinae de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Catarina, SC, Fraiburgo, 26-09-99, Borges, R.” (apple).

DIAGNOSIS. — This species is unique amongst Leucochrysa (Leucochrysa) species in several
ways. There are virtually no dark markings anywhere on the body. Amongst species found in Brazil-
ian agro-ecosystems only L. boxi has no dark markings on the abdomen, and yet it has dark longitudi-
nal stripes on the meso- and metanota. The female genitalia are also unique. The spermathecal duct is
long, yet only moderately coiled at its apex. The vela of the spermatheca is long and flattened in its api-
cal half, unlike the tubular structure of most other species. These genitalic structures are so distinctive
that we are departing from the usual procedure in describing the species based only on a female speci-
men.

The name “catarinae” refers to the state of Santa Catarina, which in English is Saint Catherine.

HEAD. — Yellow, without markings. Labial and maxillary palpi pale. Scape, pedicel and
flagellum pale.

THORAX. — Pro-, meso-, and metanota pale, without markings. Wing: Venation pale, except
first eleven costal crossveins, first three radial crossveins, middle part of Rs and last six crossveins,
gradates and marginal forks dark. Forewing length (1.81 cm): width (0.56 cm) = ratio (3.1) (Fig.
46A). Hindwing length (1.73 cm): width (0.55 cm) = ratio (3.15). Pterostigma with slightly dark pig-
mentation at basal margin.

ABDOMEN. — Green, unmarked. Female: Spermatheca apically slender, with basal half flat-
tened, coiled. Spermathecal duct long, apically coiled. Subgenitale wide with small anterio-medial
projection (Fig. 46B).

MATERIAL EXAMINED. — Known only from the holotype.

Genus Leucochrysa (Nodita) Navas, 1916

The subgenus Leucochrysa (Nodita) can be defined by the forewing triangular or ovate
intramedian cell, sinuous radial sector, and short basal extension of the female subgenitale (Brooks
and Barnard 1990). The male genitalia are inseparable from the nominate subgenus and the two
groups remain close. Species of Leuwcochrysa (Nodita) are generally smaller than Leucochrysa
(Leucochrysa) with forewing length ranging from 9 mm to about 30 mm, although a few of the larger
species, such as L. (Nodita) amazonica rival species of Leucochrysa (Leucochrysa) in size. This is the
most speciose group of chrysopids in the New World, with Brooks and Barnard (1990) listing 120 spe-
cies and several more having been subsequently described (Penny 1998, 2001). Herein we describe 24
more, previously undescribed, taxa. This subgenus will probably eventually contain more than 200
species. They are distributed from the southern U.S.A. (Penny et. al 1997) to northern Argentina
(Penny 1978). We have found 36 species in Brazilian agro-ecosystems, adults of which can be distin-
guished using the following key.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 285

KEY TO SPECIES OF LEUCOCHRYSA (NODITA) FOUND IN BRAZILIAN AGRO-ECOSYSTEMS

Scapesmithrone longitudinal stripe (Eigh8lA)s 212) 205 2s Fe ek Se Ss eee 2
Scape with two stripes (Fig. 53A), completely dark (Fig. 49A), or of other form (Fig.51A) ............. 20
Seape with dorsal longitudinal’stripe (Fig. SIA) 72s 4s. a Alas) eel ee es L. (N.) vittata
ean iaicral lonpitudinal stripe (Figi 4 7A) C40). OP 2 ee I ee BS PS eek. 3
Lateral stripe of scape well defined and wide, attaining both the anterior and posterior sides of the scape (Fig.47A) 4
Lateral stripe of scape narrow, not attaining the anterior and posterior sides of the scape (Fig.62A)......... 14
Blavelinmmpaler(Eiger/C)iecs 2 ee rect ees ears AR eh ued oly. Sratiag Seynee ty optics ecpes a 5
Elageliumicompletely (Fig: SSA) or partially dark (Fig. 47A). 2... 5.2.4 0.2 see ee ee ee te ec 7
Forewing with base of Rs, MA and inner gradate veins dark and darkly margined (Fig.61B)...... L. (N.) confusa
. Forewing with base of Rs, MA and inner gradate veins dark, but without margining or shading (Fig.47B) .... . 6
Central arch of gonarcus elongate and thin, gonocomua small (Fig. 79D, E)................ L. (N.) tenuis
. Central arch of gonarcus short and thick; gonocornua long (Fig.60E,F).................. L. (N.) barrei
Forewing with dark spots on crossveins (Fig. 70B); dark spot on the dorsal surface of the basal antennal cavity;
first two flagellar segments dark; pronotal stripes not reaching anterior margin (Fig. 70A)..... . L. (N.) maculosa
. Forewing without dark spots on crossveins (Fig. 47B); basal antennal cavity without dorsal mark; flagellum totally
dark or dark on one side; pronotal stripes reaching anterior margin (Fig. 57A).................... 8
BlaPeMuMm wit oonlyone sideldarkened (Pip. 47/A) on ces ne ae os eee ape) ese ee 9
MARC MII COMpletelyGalka(EISsI2A)) ca, 2m Gath icin hl conte as ecto 2 ay ait eines iene SATE a tees eee ee 10
Forewing with medial part of Rs and contiguous crossveins dark; marginal forks dark; hind margin of hindwing
darkened (Wie t4i//Es) eae eur Wem cree cate Akl See oe cree ners oR ae! Gre, See acta Meg AG, at SE eae L. (N.) camposi
Forewing with Rs completely pale; marginal forks pale; hind margin of hindwing pale (Fig. 68B) . . . L. (N.) interata
MONON OCIIcHlandatk spotsjonimesonotum (Big: /66A)). - 46. 4-2-4 2 e e o ee ae 2 ee 11
BINO\cicMlamdarkispots on mesonotum) (Big: SZA). 5 3. 2 oe we ne he ee ee eee, ee ee oe es es 12

. A single pair of dark spots on mesonotum (Fig. 66A); no dark spots on 2nd axillary sclerite; small dark spot on

Isteralismmaceouscape| (EIS KOOL) ewe cw Na Sp So eee Ne ava aab 6. ENG, Gy dey dhe apg daele. SWS: Gs Seale L. (N.) ictericus

. Two pair of dark spots on mesonotum; a dark spot at base of 2nd axillary sclerite (Fig. 57A); lateral surface of

Scapeicomplciciypale( HIPS SOD) eae © Grae oan, es meas Site aed ce st ee ee ae e- L. (N.) melanocera

. Lateral stripe of pronotum well defined, straight, wide, reaching the anterior and posterior margins (Fig. 58A)

39 6 0 o an Wodl enas Tel quel. CORR, eee te ata i eh a ten hae Os nr a Si a OE ie L. (N.) rodriguezi

. Lateral stripe of pronotum undulating and not reaching posterior margin; inner and outer margins of flagellum

CoUALyACaTKeENed (IPAS OFA) ease eet Aiea te MME Cite See ee NGS aes hl GERARD ont Eons eacie See aI eS 13
. Gena pale yellow; posterior margin of hindwing pale (Fig. 52B); spermatheca short, broad, with a small lateral

projection opposite juncture with spermathecal duct (Fig.52D).................... L. (N.) intermedia
. Gena red (Fig. 55C); posterior margin of hindwing darkened (Fig. 55B); spermatheca elongate and coiled

(it, SSN) oo 88s 6 Gagnon ins loro ea os oa eee ee ae tnt Peco Tome We Ee EAS Ee a Lar L.(N.) marginalis
MmGCH Awl Garkemarkinys| (MiP T4I9©) Maya cei a ruaceeee oe aaa RReM lds te Ame ae mk a ee 15
aGenarcompletely, pale (iz 1O4E) iL traces een ee eS Oy RT LIE ATES ee OQ 19
. Pronotal stripe broken into small spot at anterior margin and larger spot at mid-length (Fig. 62A); arcessus bearing

Palko mcorsalbonns| (EipO2E) weiner Vases en ee cee aes aulnee Didowion tans tr bath he de 16
. Pronotal stripe continuous, even if constricted at mid-length (Fig. 72A); arcessus not bearing pair of dorsal horns

CE eT) es tee wares eset PSS Sg VE ery. pee Lawperd His ne! io carsinteray Aaneuiiremty duaal ootecpawelarie: tered» 17
. Maxillary palps pale (Fig. 62C); dark spot on mesoscutum barely visible (Fig. 62A); dorsal horns of arcessus

projectingyposteriorly, (F1p462D) iy. easy. SSR cacks esl. Sees es Mos Be eae L. (N.) cornuta
. Maxillary palps dark apically (Fig. 48C); dark spot on mesoscutum well developed (Fig. 48C); dorsal horns of

ancessusiprojecting dorsally(Kige48E) cay 0s eich eels ses Ghote) coe UNE Gees 4 ere eau L. (N.) clepsydra
. Mesonotum bearing pair of well-developed dark spots (Fig. 72A)..................-.. L. (N.) parallela

mViesonotumicompletely pale (RigaiS Ayman ances no Chae s Has Sus Qe sitres a ee Rees le shee, Gene 18

286 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

18. Maxillary palps dark; gena completely red (Fig. 75D)... .- 2. - +2. 45-2 nee eee L. (N.) santini
18’. Maxillary palps pale; gena red only near frons (Fig. 74D). .....-.---..------+++e-ee- L. (N.) robusta
19. Gonarcus narrow, gonocornua in the form of a broad plate with small apical point (Fig. 73C) ...... L. (N.) retusa
19’. Gonarcus broad; gonocornua narrower, with more elongate point (Fig. 80E) ............... L. (N.) vignisi
20;*Scape' with two dark stripes (Fig. 53A) . «0.0. 0 se fe ee ee EE ge eG 21
20'. Dorsal surface of scape completely dark (Fig. 49A) 0 2 5 eee on cee eee eee 24
21. Pronotum(\with two pair of stripes\(Fig. 53A)).-. 1 2. ens ee ed a 22
21'_Pronotum with a single} pair of lateral'stripes|(Fig.49A)) 2.) 20 2 2s Rs 0 oo ee 23

22. Forewing with Rs and adjacent veins dark (Fig. 53B); a single pair of dark marks on abdominal tergites (Fig. 53G)
Pe ae ee ane neem. A Ie SB ek arnt Ane dS Ae Ge Bri Oo « L. (N.) lancala

22'. Forewing with most veins pale, including Rs and adjacent veins (Fig. 56B); abdomen with five pairs of dark marks
onitergites)\(Fip. SOE)... fc 0. a 5 ee eh tua Soma uh eGo cla croeetac re eo ene ne L. (N.) marquezi
23. Hindwing apically pale (Fig. 76B); vertex with lateral dark stripes and two central crescent-shaped marks (Fig. 76A);
pronotum marks confined to two pair of small spots (Fig.76A)..........-..-..-..-0-- L. (N.) scomparini
23'. Hindwing apically dark (Fig. 51B); vertex pale, except for continuation of scape stripe; pronotum with complete
darkslateralistripe(FigeSilA) 2s 20s aes ence ae eta al ee ee ee L. (N.) herioles
24. Gonocomua‘elongate, in the form of spines'((Fig.67D). 2 0.) 5 0 a 25
24/5 Gonocomua forming shorter flat plates|(Pigi65E). - = 3p wen cls) cit Sane) cele a 29
25. Forewing pterostigma weakly darkened (Fig. 67A); gonocornua twisted (Fig.67D).......... L. (N.) incognita
25’. Forewing pterostigma with well-developed dark mark (Fig. 49B); gonocornua slender, straight (Fig. SOF)... .. . 26
26), BaselofRs in the forewing darkened (Figs49B)) ps 2s) ee) ne) 27
26). Baseiof.Rs inthe forewing'pale (Fig: 54B) 0... 2. 2s eee ee ee es Sc cee ees 28
27. Flagellum dark (Fig. 52A); gonarcus strongly arched anteriorly; gonocornua forcipate; dorsal horns of arcessus
apicallyspointed (Figs49R)iam = eis 2 nso tose 2 Spares cas oe eg ee L. (N.) cruentata
27'. Flagellum pale (Fig. 68A); gonarcus only weakly arched anteriorly; gonocornua straight; dorsal horns of arcessus
apically rounded (Fig, SO EVE). a. 8 8.642. Sates Oe See ea ee on ee ee L. (N.) gossei
28. Vertex with Y-shaped dark mark behind antennal bases (Fig. 54A); a2—a3 crossvein not darkly margined; base of
forewing darkened \(Fig-S4B)i< 2a <8. 5 oo Me ew, ee ss OS ty ne L. (N.) lateralis
28’. Vertex bearing dark lateral stripes beside eyes and two medial crescent-shaped marks (Fig. 77A); a2—a3 crossvein
darklyimargined)ibut base of forewing pale (Pigad/B)) 4) ane 2 ee eee ene L. (N.) squamisetosa
29. Medial margin of pronotal stripes not straight, not well defined (Fig. 65A); abdomen bearing series of
crescent-shapedidark marks (Fig:\65€)i. 02. 6G eee ee ee eo oe ne 30
29'. Medial margin of pronotal stripes straight, well-defined (Fig. 69A); abdomen bearing longitudinal dark medial
stripe(Fig, 64@). cee ee ne ee Pe ee a SR ee ee ae 6S) ae cS ener 31]
30 Gonocomua apically directedswidestat base) (1gs055) ine ey e-news anne nein ere L. (N.) guataparensis
30’. Gonocornua apico-laterally directed, widest at mid-length (Fig. SSE) ................... L. (N.) affinis
Ses Gonocormuamuchtshortenthaniapex of arcessus|(Fig364G) i eee cme en) Gene ne eee L. (N.) furcata
3ilGonocornua’ elongate; reaching tojapex ofjarcessusi(Fig. 695) >=) 3 2. . ae ee B2
32. Gonarcus narrow (Fig. 69E); pterostigma only weakly darkened (Fig.69B)...................... 33
32’. Gonarcus broad (Fig. 63E); pterostigma heavily pigmented (Fig.63B) ........................ 34
33. Rs of forewing and adjacent veins dark; basal cavity of antennae without dorsal dark mark (Fig. 69C); apical
intersection of costal crossveins darkened; apex of gonocornua with a single point (Fig. 69E)...... L. (N.) lineata
33’. Rs of forewing pale; basal cavity of antennae with dorsal dark mark (Fig. 78C); apical intersection of costal
crossveins pale; apex of gonocomnua forked (Fig. 78E)......................... L. (N.) tabacinus

34. Complete, dark transverse band directly behind antennal base (Fig. 63D); apex of gonocornua bifurcate (Fig. 63E)

Pes cae ee oe fe okt ok ins aon, oh ere eee ah eR ATE a L. (N.) forciformis
34’. No transverse band on vertex behind antennal bases (Fig. 71D); apex of gonocornua with a single point (Fig. 71E)

Rha ei OME Choke ee SO ee Oreo ee a oe Ce men cee ono Re Geo ah oS oo oc L. (N.) michelini

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 287

Leucochrysa (Nodita) camposi Navas, 1933

DIAGNOSIS. — This species is one of several that have a dorso-lateral stripe on the scape, paired
marks on the raised medial portion of the vertex, submedial dark markings on meso- and metanota,
middle region of the Rs darkened, pale apex to the hindwing, and paired dark markings on each ab-
dominal segment. Leucochrysa (Nodita) melanocera can be separated by its pale gena, completely
dark flagellum, and pale metanotum. Leucochrysa (N.) cruentata has dark palpi and dark scape mark-
ing covering entire dorsal surface. Leucochrysa (N.) scomparini has a medial dark spot on the scape,
longitudinal lateral stripes on the vertex, and metascutellum completely dark. Leucochrysa (N.)
vittata has no dark lateral stripe on the pronotum, scape stripe is medial rather than lateral, and no dark
markings on metascutellum. Leucochrysa (N.) affinis, L. (N.) guataparensis, and L. (N.) rodriguezi
have dark antennae and much reduced dark markings on meso- and metanota. Leucochrysa (N.)
furcata, L. (N.) lineata, L. (N.) michelini, and L. (N.) tabacinus form a group of species which have
pale gena, broader and more extended lateral pronotal stripes, and dark abdominal markings com-
pletely covering tergites. However, the most distinctive feature of L. (N.) camposi is the subapical,
truncate plate of the arcessus. This feature is found in no other species encountered in
agro-ecosystems, but is found in the more northern L. (N.) amazonica. Both species have other charac-
teristics in common, such as large size (forewing length > 20 mm), crescent-shaped markings on the
vertex, and dark Rs vein at mid-length. However, L. (N.) amazonica bears no scape stripe, nor does it
have dark metanotal and abdominal tergal markings.

HEAD. — Yellow. Frons without marks. Gena with thin red stripe. Maxillary and labial palpi pale
(Fig. 47A, C). Vertex yellow, raised, marked with two medial, crescent-shaped red bands. Antennal
scape and pedicel with dark red, dorso-lateral stripe continued onto antennal fossa; flagellum basally
dark on lateral margin, apically pale.

THORAX. — Green with yellow dorsal median stripe. Pronotum with anterio-lateral darkened
band (Fig. 47A). Mesonotum marked with red spots on prescutum and scutum. Metanotum with red
spots on scutum and scutellum (Fig. 47G). Wings: Venation green, except marginal endings of costal
crossveins, middle part of Rs, Psm-Psc crossveins 1—7, forks of marginal veins and anal veins dark.
Pterostigma slightly darkened basally. Length (2.08—2.15 cm); width (0.64—0.76 cm) (Fig. 47B).
Hindwing pterostigma well marked. Hindwing green, except last radial crossvein, last costal
crossvein, middle part of Rs, and posterior marginal area darkened (Fig. 47B). Length (1.78 cm):
width (0.59 cm) = ratio (3.02).

ABDOMEN. — Green, with red marks on all tergites (Fig. 471). Male: Sternite 8 + 9 not com-
pletely fused; microtholi present (Fig. 47H). Gonarcus thick with triangular lateral arms. Gonocornua
long and apically decurved. Entoprocessus absent. Arcessus with subapical dorsal plate and apical
ventrally-folded plate with medial point (Fig. 47D, E, F). Female: Subgenitale with short,
anterio-medial projection (Fig. 47J). Bursa sac-like, folded at apex, with pair of filamentous, multi-
ply-forked accessory glands. Spermatheca long, ventral impression short.

MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 18 July 1998, Freitas, S. (1)(rub-
ber)(CAS); Sao Paulo: Taquaritinga, 8 June 1993, Xavier, A. L. Q. (1“)(orange); 21 October 1992,
Freitas, S. (1 2)(orange); Catanduva, May 1994, Silva, J. L. (1%)(orange); Taquaral, 13 March 1991,
Freitas, S. (1o*)(orange); 5 March1991, Freitas, S. (1 2)(orange); Jaquariuna, May 1992, Kubo, R. K.
(50°)(orange); Jaboticabal, 26 August 1995, Freitas, S.(1°)(guava); Taiuva, 27 May 1992, Pessoa, R.
(1o)(orange); 8 June 1990, Freitas, S. (12 )(orange); 11 September 1995, Ferreira, R. J. (22 )(guava);
21 October 1995, Ferreira, R. J. (1 2)(guava); October 1989, Fernandes, O. D. (12); 10 July 1998,
Freitas, S. (1 2)(CAS).

288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

Leucochrysa (Nodita) clepsydra Banks, 1918

DIAGNOSIS. — Leucochrysa (N.) clepsydra is associated with L. (N.) cornuta by the thin, red
stripe of the gena; pale antennae with thin, incomplete dorsal stripe of the scape; pair of dark lateral
spots on either side of pronotum of unequal size; pair of dark mesonotal spots; forewing Rs with dark
area at mid-length, and pale apex of hindwing. Differences between the two species include dark
maxillary palpi; two small dark spots on vertex; small gonarcus with vertically elongate lateral arms
and longer pair of subapical dorsal horns in L. (N.) clepsydra.

HEAD. — Yellowish green. Gena with reddish stripe near frons. Apical maxillary palpimeres
dark. Vertex marked with two, diverging, elongate, red spots (Fig. 48A,C). Antennae pale with short
red stripe dorso-laterally on scape.

THORAX. — Cervical sclerites red. Pronotum with two red spots laterally on either side.
Mesonotum with pair of red spots (Fig.48A). Metanotum unmarked. Wings: Forewing venation
green, except both junctures of costal crossveins, apex of Rs, gradates and marginal forks dark (Fig.
48B). Pterostigma weakly darkened basally. Forewing length (1.46 cm): width (0.50 cm) = ratio
(2292):

ABDOMEN. — Green, marked with red. Male: Gonarcus short, poorly defined, with small, ovate
lateral arms. Gonocornua well developed, apically decurved, surface with tiny setae. Arcessus poorly
sclerotized, with pair of large, dorsal, subapical horns between gonocornua and apico-medial
point and two, small, rounded, lateral lobes. Gonossacus with few, scattered, lateral gonosetae (Fig.
48D, E, F).

MATERIAL EXAMINED. — Colombia: Caldas, May, 4900 ft. [1490 m], Fassl, four syntypes (de-
posited in Museum of Comparative Zoology, Harvard University), Brazil: Sao Paulo: Taquaritinga,
8 January 1993, Xavier, A. L. Q. (1o)(orange).

Leucochrysa (Nodita) cruentata (Schneider, 1851)

DIAGNOSIS. — This species is part of the complex having red gena, crescent-shaped marks on the
vertex, dark submedial markings on meso- and metanotum, and extensive dark markings on abdomi-
nal tergites. However, the combination of dark palpi, completely red dorsal surface of the scape, shad-
ing of the origin of Rs on the forewing, and large, incurved gonocornua will separate this from all other
species.

HEAD. — Green. Gena red. Frons unmarked. Clypeus laterally marked with red (Fig. 49C).
Maxillary and labial palpimeres dark. Vertex with divergent red stripes on medial raised area. Scape
completely red dorsally, the red coloration extends onto basal membrane, pale ventrally; pedicel with
red ring not shown on illustration; flagellum pale (Fig. 49A).

THORAX. — Green, marked with red. Pronotum as wide as long, with lateral red stripes, broader
at mid-length and converging anteriorly. Meso- and metanotum with variegated red marks (Fig. 49A).
Wings: Forewing venation pale, except costal crossveins, R-Rs crossveins, Rs-Psm, inner and outer
gradates, marginal forks, and Rs dark. Origin of Rs with brown shading. Wing base with linear red
mark (Fig. 49B). Pterostigma with basal dark spot. Length (2.50 cm): width (0.98 cm) = ratio (2.55).
Hindwing venation pale, except costal crossveins, and inner and outer gradates dark (Fig. 49B).
Pterostigma with heavy dark mark basally. Length (2.13 cm): width (0.73 cm) = ratio (2.92).

ABDOMEN. — Tergites 3, 5, 6, 7, and 8 with reddish brown crescent-shaped marks (Fig. 49D).
Male: Microtholi absent. Lateral arms of gonarcus ovate. Gonocornua large, widely separated ba-
sally, apically incurved at a sharp angle. Arcessus broad; dorsal surface poorly sclerotized, with pair
of subapical, dorsal horns, apical portion decurved, with medial hook and membranous, rounded lat-
eral lobes (Fig. 49E, F, G, H). Female: Subgenitale with apical sclerotized sac with many gonocristae

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 289

(Fig. 49J). Spermatheca large, thick. Vela and bursal duct well developed. Spermathecal duct thin,
with apical brush (Fig. 491).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 21 December 1993, Freitas, S.,
(1o)(rubber); 17 August 1995, Freitas, S.(2o°l 2 (orange); 11 October 1995, Freitas, S. (1 )(rubber);
10 February 1998, Freitas, S. (1 )(guava); 5 July 1998, Freitas, S.(12)(CAS); 5 August 1998, Freitas,
S. (1 2)(guava); 10 March 1999, Freitas, S. (1 2)(CAS).

Leucochrysa (Nodita) gossei (Kimmins, 1940)

DIAGNOSIS. — This species appears to be closely related to L. (N.) cruentata because of the com-
pletely red dorsum of the scape, red gena, dark lateral markings of both meso- and metanota, dark
shading of the origin of Rs on the forewing. However, L. (N.) gossei has pale palpi and the tergal marks
on the abdomen are quite different. In the male genitalia, the gonocornua of L. (N.) gossei are slightly
turned outward, while in L. (N.) crentata they are strongly medially curved.

HEAD. — Yellow. Gena red. Apical maxillary palpimeres dark. Vertex with pair of small, thin
red marks. Scape completely red dorsally, pale ventrally; pedicel pale; flagellum pale (Fig. 50A, C).

THORAX. — Green. Pronotum longer than wide, with lateral stripes dark red. Meso- and
metanotum yellow with brown spots (Fig. 50A). Wings: Forewing venation green, except gradates,
R-Rs crossveins, apices of Rs-inner gradates, marginal forked and unforked veins, and Psm-Psc
crossveins dark. Gradates not darkly bordered. Pterostigma with dark basal mark. Length (1.47 cm):
width (0.56 cm) = ratio (2.63). Hindwing with acute apex. Pterostigmal dark spot well defined. Vena-
tion green, except apex of Rs, gradates, and marginal forks dark (Fig. 50B). Length (1.27 cm): width
(0.41 cm) = ratio (3.10).

ABDOMEN. — Yellow with large dark spot on tergite 3, two small ones on tergite 4, large one on
tergite 6, and two small dark marks on tergite 7 (Fig. 50D). Male: Dorsal apodeme of ectoproct forked
near callus cerci. Microtholi absent. Gonarcus arcuate; lateral arms short. Gonocornua elongate, par-
allel and apically decurved. Arcessus broad; with pair of dorsal plates elongate, upturned, apically
truncate; apex with small medial horn and acute lateral lobes. Gonosaccus with few gonosetae (Fig.
50E, F, G). Female: Spermatheca twice-recurved. Bursa wrinkled with a pair of long accessory
glands. Subgenitale with short, rounded lateral lobes and elongate medial projection (Fig. 50H, 1).

MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 14 September 98, Freitas, S. (1 *)(rub-
ber); SAo Paulo: Luis Antonio, 9 August 1993, Freitas, S. (12)(CAS); 8 June 1994, Freitas, S.
(1%)(CAS).

REMARKS. — This species was originally described by Walker (1853) as Chrysopa conformis.
However, Walker’s name is an objective junior homonym of Hemerobius conformis Rambur, 1842, a
name which was subsequently transferred to the genus Chrysopa by Walker himself in 1853.

Leucochrysa (Nodita) heriocles Banks, 1944

DIAGNOSIS. — Leucochrysa (Nodita) heriocles is distinctive in being the only species with a
dark apex to the hindwing found in the agro-ecosystem. Several other species found further north have
this trait. Of these species only L. (N.) apicata, L. (N.) indiga, and L. (N.) postica have dark palpi, api-
cally expanded stripe on the scape, and lateral red pronotal stripe (see Penny 2001). However, L. (N.)
indiga has a complete dark band across the frons and L. (N.) postica has an almost complete band. Nei-
ther of these species have dark markings on meso- and metanotum. Leucochrysa (N.) apicata and L.
(N.) heriocles are close, and may in fact be the same species, both having elongate gonocornua which
approximate or cross apically, but L. (N.) heriocles males have a distinctive scalloping and apical
point to sternite 9, which is not nearly so accentuated in L. (N.) apicata.

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HEAD. — Yellow. Narrow red band across clypeus under and between anterior tentorial pits.
Gena red. Apical labial and maxillary palpi dark. Vertex pale, unmarked. Scape with red stripe
dorso-laterally expanded to complete ring apically, basally extended onto basal membrane; pedicel
with apical dark ring; flagellum pale, except for dark stripe along medial margin of antennomeres |—3
(Fig. 51A, C).

THORAX. — Yellowish green. Pronotum longer than wide with lateral dark red stripe.
Mesonotum with diffuse reddish marks, especially pronounced along anterior margin. Metanotum
pale, unmarked (Fig. 51 A). Wings: Forewing longitudinal and some associated crossveins dark. Dark
pterostigmal spot quite evident. Gradates, Psm—Psc crossveins; and forked marginal veins bordered
(Fig. 51B). Length (1.18 cm): width (0.42 cm) = ratio (2.81). Hindwing acute; apex with diffuse dark
spot. Dark pterostigmal spot well defined. Rs, costal crossveins black; outer gradates, and forked mar-
ginal veins dark. Outer gradates darkly margined (Fig. 51B). Length (0.97 cm): width (0.30 cm) = ra-
tio (3.23).

ABDOMEN. — Green with two small dark spots on tergite | and other larger ones covering ter-
gites 3 and 6 (Fig. 51H). Male: Sternites 2-8 with dense microtholi. Erect setae on apical sternite
stouter than on tergites and arising from pit in cuticle. Apex of sternite 9 scalloped laterally, forming
medial point lined dorsally by band of gonocristae (Fig. 51 I, J). Gonarcus arcuate, with short, ovate
lateral arms. Gonocornua elongate, medially angled, apically approximated. Arcessus broad basally,
with pair of small, dorsal, subapical horns and small apico-medial hook flanked by two small rounded
lobes. Gonosaccus with sparse gonosetae. Female: Spermatheca elongate, distally coiled (Fig. 51G).

MATERIAL EXAMINED. — Surinam: Paramaribo, Charlesburg, 5 May 1941, Geijskes, syntype
male (deposited in Museum of Comparative Zoology, Harvard University), Brazil: Mato Grosso,
Itiquira, 23 March 97 (1 ")(rubber); 18 February 1996, Scomparin, C. H. J. (1o)(rubber); 10 Novem-
ber 97, Scomparin, C. H. J. (30°42)(rubber).

Leucochrysa (Nodita) intermedia (Schneider, 1851)

DIAGNOSIS. — The combination of dark antennae, lack of dark markings on meso- and
metanotum, and dark markings only on tergites | and 2 will separate this species from all others. The
lateral projection of the spermatheca vela and tiny pits on the accessory glands are also distinctive for
L. (Nodita) intermedia. The closest species is probably L. (N.) marginalis, from which L. (N.)
intermedia can be separated by its larger size (forewing length 19 mm) and pale posterior margin of
the hindwing.

HEAD. — Yellow, somewhat red-tinged on clypeus, frons and vertex. Vertex with V-shaped me-
dial dark mark. Scape with red stripe dorso-laterally; pedicel dark-ringed subapically; flagellum dark
(Fig.52A).

THORAX. — Pronotum yellowish green with pale red stripe laterally. Meso- and metanotum pale,
unmarked. Wing: Fore- and hindwing pterostigma heavily pigmented basally. Venation pale, except
origin of Rs, costal crossveins, middle part of Rs and contiguous veins, inner gradates, pseudocubital
crossveins and branches of forked marginal veins dark. Length (1.89 cm): width (0.70 cm) = ratio
(2.70). Hindwing venation pale, except middle costal crossveins, Rs-inner gradate crossveins and
contiguous veins dark (Fig. 52B). Length (1.71 cm): width (0.54 cm) = ratio (3.17).

ABDOMEN. — Green. Tergites | and 2 with dark red markings (Fig. 52C). Female: Bursa cop-
ulatrix sac-like with pair of filamentous accessory glands bearing tiny, circular, sclerotized pits on the
surface (Fig. 52D). Subgenitale two short lateral lobes and large anterio-medial projection (Fig. 52E).
Spermatheca short, thick; vela with small lateral projection (Fig. 52D).

MATERIAL EXAMINED. — Brazil: Santa Catarina: Sao Joaquim, March 1999, Borges, R.
(1 2)(apple).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 291

Leucochrysa (Nodita) lancala Banks, 1944

DIAGNOSIS. — The most distinctive characteristic of this species is the bird’s-head-shaped api-
cal projection of the arcessus. This species appears to be most closely related to L. (N.) guataparensis
which also has the distinctively shaped apex of the arcessus, but the dark palpi, unconnected red
stripes on the scape, and thinner gonocornua sets L. (N.) /ancala apart.

HEAD. — Pale yellow. Frons with small, reddish brown spot between and below antennae. Third
and fourth maxillary palpimeres and basal half of fifth segment dark. Gena red. Vertex yellow with
thin, red, medial, V-shaped mark; on raised medial area two faint, widely-spaced, red, cres-
cent-shaped marks. Scape pale with dorsolateral red stripe, continued onto antennal base and short
dorso-medial red spot; pedicel pale with dark subapical ring; flagellum pale with dark medial stripe
basally (Fig. 53A, C).

THORAX. — Cervical sclerites red. Pronotum yellow medially; green submedially; red stripe on
lateral margin, tapering posteriorly; posterio-medial half with thin black stripe. Mesonotum pale with
red tinge, more pronounced along anterior margin. Metanotum pale with red stripe laterally (Fig.
53A). Wing: Fore- and hindwing pterostigma dark. Venation green, except middle costal crossveins,
origin of Rs, fourth Rs and contiguous veins, gradates, marginal forks and posterior ends of outer
pseudocutibal veins 4—S dark. Eight inner and nine outer gradate veins. Length (1.72—1.86 cm); width
(0.55—0.67cm). Hindwing apically acute. Venation pale, except middle of Rs and contiguous
crossveins, gradates, and marginal forks dark. Length (1.55—1.74 cm); width (0.44—0.53 cm) (Fig.
DS):

ABDOMEN. — Green; mid-line yellow (Fig. 53G). Male: Apex of ectoproct truncated obliquely.
Anterior half of sternite 8 + 9 with microtholi and posterior half with dense, short, black setae.
Gonarcus thick, short, with ovate lateral arms. Gonocornua long, tapering, dorsally ridged, nearly
straight, but with slight apical twist, and somewhat angled medially. Arcessus narrow, bilobed
apodeme basally, strongly arched apically, with well-sclerotized apico-medial hook without lateral
lobes (Fig. 53D, E, F). Female: Spermatheca thick, sinuous; ventral impression short. Bursa convo-
luted. Subgenitale with swollen lateral lobes forming medial groove; anterio-medial projection finely
rugose (Fig. 53H, I, J).

MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 19 April 1993, Freitas, S. (1)(or-
ange); 10 September 1999, Freitas, S. (1 2); 10 October 1995, Freitas, S. (1o)(orange); March 1996,
Freitas, S. (1 2)(guava)(CAS); 6 February 1997, Freitas, S. (12); March 1996, Freitas, S. (12);
Birigui, 21 February 1996, Scomparin, C. H. J. (12)(corn); Mato Grosso: Itiquira, 18 September
1996, (3220)(rubber); 18 November 1996, Scomparin, C. H. J. (12)(CAS); 30 December 1996,
Scomparin, C. H. J. (1o)(rubber); 23 January 1997, Scomparin, C. H. J. (1)(rubber).

Leucochrysa (Nodita) lateralis Navas, 1913c

DIAGNOSIS. — This is one of several species with completely dark dorsal surface of the scape, V-
or Y-shaped dark mark behind antennae on vertex, and dark markings on the meso- and metanotum.
Of these species, L. (N.) gossei has dark maxillary palpi; L. (N.) cruentata has forewing shading at the
origin of Rs; and L. (N.) cruentata and L. (N.) affinis have much larger lateral stripes on the pronotum.
Perhaps the most closely related species is the more northern L. (N.) virginiae, which also has a pale
pronotum, although with a small red spot in the anterio-lateral corner (Penny 1998). However, L. (N.)
virginiae has a complete, broad dark band across the frons, which is not present in L. (N.) lateralis.
The most distinctive feature of this species is the lateral projection of the bursal vela.

HEAD. — Yellow with red markings. Gena red. Clypeus pale with slight red tinge. Maxillary and
labial palpi pale. Frons pale with short red bands below antennal bases and small medial red mark be-

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tween antennal bases. Anterior part of vertex with Y-shaped mark (Fig. 54C). Scape dark brown
throughout; pedicel pale with subapical dark ring (Fig. 54A); flagellum pale.

THORAX. — Pronotum green with pair of small red spots on anterio-lateral corners. Anterior
margin of mesoprescutum and mesoscutum laterally reddish brown (Fig. 54A). Wing: Pterostigma
with slight indication of dark mark basally. Forewing base and second axillary sclerite with brownish
red spot. Venation pale, except ends of first eight costal crossveins, all of remaining costal crossveins;
radial sector crossveins, gradates, Psm-Psc crossveins, and forked and unforked marginal veins dark
(Fig.54B). Forewing length (1.73 cm): width (0.59 cm) = ratio 2.93. Hindwing venation pale, except
costal crossveins and gradates dark. Length (1.45 cm): width (0.44) = ratio 3.30.

ABDOMEN. — Green, with reddish brown marks on tergites 2, 3, 6, and 7 (Fig. 54A). Female:
Spermatheca short, pill-box-shaped; vela with lateral projection. Spermathecal duct moderately long,
twisted but not coiled, becoming wider near spermatheca. (Fig. 54D). Subgenitale broad, without me-
dial projection (Fig. 54E).

MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 18 October 1996, Scomparin, C. H. J.
(1 2)(rubber).

Leucochrysa (Nodita) marginalis Banks, 1915

DIAGNOSIS. — This is one of a group of species with pale palpi, red gena, dorso-lateral stripe on
the scape, pair of crescent-shaped marks on the vertex, dark lateral stripes on the pronotum, and dark-
ened area in the middle of the Rs of the forewing. However, only a few other species in the group have
dark antennae: L. (N.) intermedia, L. (N.) lineata, L. (N.) melanocera, L. (N.) michelini, L. (N.)
rodriguezi, and L. (N.) tabacinus. Of these species, all have abdominal markings and more extensive
meso- and metanotal markings than L. (N.) marginalis. Males of none of the other species have the
subapical truncate plate of the arcessus of L. (N.) marginalis.

HEAD. — Yellow. Frons with tiny red spot below and between antennae. Gena red. Maxillary
and labial palpi pale. Vertex with pair of medial, short, red, crescent-shaped stripes. Scape with red
dorso-lateral stripe; pedicel pale with subapical dark ring; first 17 segments of flagellum black and
more apical segments fuscous (Fig. 55A, C).

THORAX. — Pronotum pale with lateral dark stripe, which is restricted at mid-length (Fig. 55A).
Mescutum pale with pair of small red spots. Metanotum pale, unmarked. Wings: Forewing venation
green, except marginal junctions of costal crossveins, origin of Rs, and middle part of Rs and contigu-
ous crossveins dark. Gradates pale. Length (1.63 cm): width (0.59 cm) = ratio (2.76). Hindwing vena-
tion green, except endings of basal costal crossveins, middle part of Rs and contiguous veins, and
branched endings of forked marginal veins dark. Shading along hind margin at forked marginal veins.
Length (1.41 cm): width (0.45 cm) = ratio (3.13). Fore- and hindwing pterostigma dark basally (Fig.
55B).

ABDOMEN. — Tergites without dark markings. Male: Sternites 8 and 9 not fused. Microtholi
present. Dorsal apodeme of ectoproct thick, well sclerotized near callus cerci (Fig. 55D). Gonarcus
thick, arched; lateral arms well developed, triangular. Gonocornua elongate but not reaching apex of
arcessus, thick at base. Arcessus large with subapical medial plate and small, depressed apico-medial
hook (Fig. 55E, F, G). Gonosaccus with few gonosetae, continued as second sac-like structure above
sternite 9. Female: Spermatheca short with short spermathecal duct; ventral impression short. Bursa
sac-like with pair of filamentous accessory glands (Fig. 551). Subgenitale rounded with small median
projection (Fig. 55H).

MATERIAL EXAMINED. — Bolivia: Rio Longo, 750 m, Fassl, holotype male (deposited in Mu-
seum of Comparative Zoology, Harvard University), Brazil: Sao Paulo: Balsamo, 15-X-97, Berg-
man, E. (121o)(rubber).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 293

Leucochrysa (Nodita) marquezi Navas, 1913a

DIAGNOSIS. — The oblique dark stripes at the posterior margin of the pronotum link L. (N.)
marquezi with L. (N.) affinis, L. (N.) guataparensis, L. (N.) internata, and L. (N.) lancala. Of these
species, L. (N.) marquezi is probably most closely related to L. (N.) lancala, with which it shares dark
palpi and a dark mark on the medial surface of the scape. They also share sigmoidally curved
gonocornua and quadrate lateral arms of the gonarcus of the male genitalia. However, L. (N.) lancala
has more extensive dark markings on the meso- and metanotum, no dark markings on abdominal ter-
gites 2 and 3, and a distinctively curved, bird’s head shape to the apex of the arcessus (Fig. 53D).

HEAD. — Yellow. Frons with small reddish brown spot between and below antennae (Fig. 56C).
Gena red (Fig. 56D). Apical maxillary palpimeres dark. Vertex pale with red, V-shaped mark behind
antennal bases; posteriorly pair of crescent-shaped red stripes on raised medial area (Fig. 56A, C).
Scape yellow with dorso-lateral red stripe continued onto antennal fossa, and short dorso-lateral red
mark along medial margin (Fig. 56A); pedicel pale with subapical dark ring; flagellum pale.

THORAX. — Cervical sclerites red (Fig. 56C, D). Pronotum yellow medially, green submedially,
with red stripe on lateral margin which tapers toward posterior margin; postero-medial region with
pair of oblique, curved, black stripes about halfway between center line and margin. Mesonotum yel-
low with dark marks on prescutum and scutum (Fig. 56A). Wings: Forewing pterostigma with
slightly dark pigmentation basally. Venation green, except first, second and last radial crossvein, apex
of Rs, Rs-Psm crossveins 2—4; Psm-Psc crossveins, gradates, marginal forked and unforked veins
dark (Fig. 56B). Forewing length (1.8—2.1 cm); width (0.64—0.72 cm). Hindwing acute. Pterostigma
with well-developed basal dark mark. Venation green, except apex of Rs, gradates, marginal forked
branches and apical costal crossveins dark (Fig. 56B). Length (1.44 cm): width (0.46 cm) = ratio
(3.13).

ABDOMEN. — Green with yellow mid-line and red marks on tergites 2, 3, 6 and 7 (Fig. 56E).
Male: Ectoproct apically truncate. Sternites 8 and 9 not fused; with many microtholi. Gonarcus thick,
with vertically quadrate lateral arms. Gonocornua long, slightly sinuous, medially slanted. Arcessus
short, double apodemes basally and with strong apical hook without small lateral lobes (Fig. 56H, I).
Female: Spermathecae curved, ventral impression short. Bursa convoluted with pair of filamentous
accessory glands (Fig. 56F, G). Subgenitale sclerotized, anterio-medial part elevated above mem-
brane.

MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 10 February 1998, Freitas, S. (171 2);
10 March 1999, Freitas, S. (1); Mato Grosso: Itiquira, 20 January 1997, Freitas, S. (30°l 2 )(rub-
ber)(SDF/CAS); 23 January 1997, Freitas, S. (1o°1?%)(rubber); 13 September 1998, Freitas, S.
(12)(CAS).

Leucochrysa (Nodita) melanocera Navas, 1916

DIAGNOSIS. — This species is most notable for the lack of markings—both on the face and
metanotum. Also, abdominal markings, although present on all segments, are quite small. The distinc-
tive spotting pattern of the mesonotum and second axillary sclerite indicate a fairly close relationship
with L. (N.) barrei and L. (N.) parallela, but L. (N.) barrei has dark palpi and gena, and both have pale
antennae. The pentagonal-shaped subgenitale and distinctly two-parted spermathecal vela can also
help distinguish this species.

HEAD. — Yellow. Frons, clypeus and gena without dark markings. Maxillary and labial palpi
pale. Vertex with two dark marks anteriorly and weak red marks on raised medial area. Scape and
pedicel with broad, dorso-lateral dark stripes; flagellum dark.

THORAX. — Pronotum greenish yellow with thin, lateral, red stripes widening at apices.
Mesoprescutum with black spots laterally; mesoscutum with pair of dark spots medially. Second

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axillary sclerite with dark spot (Fig. 57A). Wings: Pterostigma with faint dark spot basally. Forewing
venation pale with costal crossvein endings, origin of radial sector, first Rs-Psm crossveins, middle
part of Rs and adjacent veins, gradates, and forked marginal veins dark. Length (1.77 cm): width
(0.65 cm) = ratio (2.72). Hindwing acute. Venation pale, except costal crossveins, middle part of Rs
and some marginal forks dark (Fig. 57B). Length (1.52 cm): width (0.50 cm) = ratio (3.04).

ABDOMEN. — Yellowish green. Tergites with lateral dark red spots (Fig. 57A). Female:
Spermatheca with vela divided into two parts (Fig. 57C). Bursa with pair of filamentous accessory
glands. Subgenitale pentangular (Fig. 57D).

MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 18 February 1996, Freitas, S.
(1 2)(rubber).

Leucochrysa (Nodita) rodriguezi (Navas, 1913a)

DIAGNOSIS. — This is one of a large group of species with red gena, dorso-lateral stripes on the
scape, crescent-shaped marks on the vertex, dark red lateral stripes on the pronotum, and dark area at
mid-length of forewing Rs. It is one of six species of this group with dark antennae found in Brazilian
agro-ecosystems. Of these species, only L. (N.) marginalis has similar small linear dark markings on
the mesoscutum. However, L. (N.) marginalis has no abdominal tergite markings and has a distinctive
subapical, truncate plate on the arcessus. The gonocornua with dorso-basal tooth is a distinctive fea-
ture of L. (N.) rodriguezi found in no other species.

HEAD. — Yellow with red markings. Vertex with pair of oblique, crescent-shaped red marks on
medial raised area. Small red spot between antennal bases. Gena red. Maxillary and labial palpi pale.
Scape with dark red dorso-lateral stripe; pedicel with small dorso-lateral dark spot; flagellum black,
basal antennomeres laterally more intensively black (Fig. 58A, D).

THORAX. — Yellowish green. Pronotum with lateral red stripes. Mesoscutum pale with pair of
medial red spots and premesoscutum with pair of lateral, diffuse, pale red spots. Metanotum pale,
without marks (Fig. 58A). Pterostigma with dark basal mark. Forewing venation green, except costal
crossveins 5—15, origin of Rs, Rs-Psm crossveins 1—6, apex of Rs, inner gradates 5—7 and outer
gradates 7-9, and Psm-Psc crossveins dark. Length (16.9—17.8 mm); width (5.3—6.4 mm). Hindwing
venation pale, except costal crossveins 7—10, and outer gradates dark Fig. 58B). Length (1.44 cm):
width (0.47 cm) = ratio (3.06).

ABDOMEN. — Yellow with pair of small red stripes on each tergite (Fig. 58C). Male: Apex of
ectoproct with several long setae. Sternites 8 + 9 fused and apically bilobed; with many microtholi
(Fig. S8E). Gonarcus thick, only slightly arched; lateral arms short, ovate. Gonocornua elongate,
forcipate with dorso-basal tooth. Arcessus with dorso-medial, subapical, acute projection and ven-
trally decurved apical hook flanked by small, rounded lateral lobes. Gonosaccus with few scattered
setae (Fig. 58G, H). Female: Cavity between subgenitale and sternite 8 large and sclerotized, with
tiny microtrichia. Subgenitale apically bilobed, with anterio-medial projection. Spermatheca short
with ventral impression short. Apex of spermathecal duct strongly dilated (Fig. 581).

MATERIAL EXAMINED. — Brazil: S40 Paulo — Jaboticabal, 27 August 1996, Domenici, M. G.
(1); 29 August 1995, Ferreira, R. J. (1o°)(CAS); 3 November 1989, Jacob, M. (1 )(orange); 9 Octo-
ber 1989, Carmo, O.B. (1 2); 23 March 1996, Fernandes, O. A. (1 2); 9 August 1996, Freitas, S. (10);
Taquaritinga, 4 September 1993, Xavier, A. L. Q. (1 2)(orange); 4 September 1993, Narciso, R.
(1o°)(orange); Birigui, 21 February 1996, Scomparin, C. H. J. (1o)(corn); Mato Grosso: Itiquira, 9
November 1999, Freitas, S. (1o)(rubber); 10 September 1996, Scomparin, C. H. J. (1 2)(rubber); 14
December 1996, Scomparin, C. H. J. (1 2)(rubber); Bahia: Juazeiro, 6 March 1996, Ferreira, R. J.
(2)(cashew nut).

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 295

Leucochrysa (Nodita) affinis de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Sao Paulo: Jaboticabal, 5 June 1995, Freitas, S.” (orange).

DIAGNOSIS. — This is a member of a group of species with the dorsal surface of the scape com-
pletely red, while the ventral surface is completely pale. All species in this group appear to have at
least some dark markings on the mesonotum. Leucochrysa (N.) affinis appears intermediate with the
group of species having oblique dark marks at the posterior part of the pronotum, and is the only spe-
cies with completely red dorsal scape to have oblique dark marks on the pronotum. A similar species 1s
L. (N.) cruentata, which has the dark dorsal side of the scape, dark flagellum, and extensive segmental
markings on the abdominal tergites. It also has large, angled gonocornua, but they are forcipate and
rather evenly tapered in L. (N.) cruentata, whereas the gonocornua of L. (N.) affinis are distinctively
flattened, widened at mid-length, and apically divergent. Leucochrysa (N.) affinis actually appears
most closely related to L. (N.) guataparensis, which does not have a completely red dorsal surface on
the scape, but has markings more like two longitudinal stripes partially fused along the mid-line. The
male gonocornua of both species are highly flattened, but in L. (N.) guataparensis they are triangular,
while in L. (N.) affinis they are strongly elbowed. Leucochrysa (N.) affinis also has dark antennae,
which are pale in L. (N.) guataparensis.

The name affinis is Latin for “related to” and refers to the close relationship of this species to L.
(N.) guataparensis.

HEAD. — Yellow. Gena red. Vertex with V-shaped red mark behind antennal bases. Maxillary
and labial palpi pale. Scape red dorsally, pale ventrally; pedicel and flagellum black (Fig. 59A,D).

THORAX. — Pronotum pale green with lateral red stripe, fused at posterio-lateral margin with
second, oblique, thin, red mark on posterior half (Fig. 59A). Meso and metanotum pale, unmarked.
Wings: Forewing pterostigma with faint dark mark basally. Venation green; except costal crossveins,
R-Rs crossveins, gradates, and apices of marginal forks dark. Length (1.76 cm): width (0.64 cm) =ra-
tio (2.75). Hindwing pterostigma with well-developed dark mark basally. Venation pale, except mid-
dle part of Rs and contiguous veins, gradates, and apices of marginal forks dark (Fig. 59B). Length
(1.47 cm): width (0.48 cm) = ratio (3.06).

ABDOMEN. — Green with red marks (Fig. 59C). Male: Microtholi on sternite 8 + 9. Gonarcus
short and broad, with ovate lateral arms. Gonocornua dorso-ventrally flattened and broadly expanded
at mid-length, apically pointed with divergent apices. Arcessus basally broad, without dorsal horns;
decurved apical hook without well-developed lateral lobes (Fig. 59E,F). Female: Spermatheca short;
ventral impression short. Bursa convoluted with pair of filamentous accessory glands.

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 4 December 1996, Freitas, S.
(3c paratypes)(orange)(SDF/CAS); Mato Grosso: Itiquira, 5 April 1996, Scomparin, C. H. J. (1¢
paratype)(rubber); 20 January 1997, Freitas, S. (12 paratype)(CAS); 16 August 1997, Freitas, S.
(8022 paratypes)(rubber).

Leucochrysa (Nodita) barrei de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 2-IX-97, Scomparin, C. H., SP79 o” (rubber).

DIAGNOSIS. — This is one of the species with well-defined dark spots on the mesoscutum. Of
these species, L. (N.) clepsydra has a small spot rather than dorso-lateral stripe on the scape, dark red
spots rather than lateral stripe on pronotum, and prominent dorsal horns of the arcessus. Unlike L. (N.)
barrei, L. (N.) ictericus has pale palpi, no second axillary sclerite markings, and basally and apically
toothed gonocornua. Leucochrysa (N.) melanocera has pale palpi, thin lateral pronotal stripes, and

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small dark markings on all abdominal tergites. The highly arched form of the arcessus and
gonocornua, as well as small ventral lobe at mid-length of the gonocornua are unique to L. (N.) barrei.

This species is dedicated to Lionel Barré, who enthusiastically initiated the use of chrysopids for
biological control of rubber tree pests in Brazil.

HEAD. — Yellow. Frons and clypeus pale, unmarked. Gena with small red stripe near frons.
Maxillary palpi basally pale, dark in middle, and apical half of apical segment pale. Labial palpi pale
(Fig. 60C, D). Vertex yellow, unmarked. Scape with dorsolateral red stripe; pedicel with apical dark
ring; flagellum pale.

THORAX. — Pronotum pale with irregular, lateral, thin, dark red stripe. Mesonotum with pair of
dark circular spots submedially. Mesonotal second axillary sclerite red basally. Metanotum pale, un-
marked (Fig. 60A). Wing: Forewing pterostigma faintly darkened basally. Venation green, except
costal, radial and Psm-Psc crossveins, gradates, and apices of marginal forks dark. Length (2.75 cm):
width (1.0 cm) = ratio (2.75). Hindwing pterostigma with well-developed dark basal spot. Venation
green, except apex of Rs and outer gradates dark (Fig. 60B). Length (2.30 cm): width (0.75 cm) = ratio
(3.07).

ABDOMEN. — Green with dark marks on tergites 3 and 4. Male: Abdominal sternites without
microtholi. Gonarcus short, with no vertical arch; lateral arms ovate, longer than high (Fig. 60E).
Gonocornua slender, forcipate and decurved, with small ventral lobe at mid-length (Fig. 60F).
Arcessus strongly arched high above gonocornua, with subapical lateral ridge and decurved apical
hook. Gonosaccus without gonosetae.

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) confusa de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Fraiburgo, SC, 26-IX-99, Maga, SP83, Borges R.” (apple).

DIAGNOSIS. — This is one of four species in the region which have the origin of the radial sector
heavily shaded. Two of these, L. (N.) cruentata and L. (N.) gossei, have darkened dorsal surface of the
scape and belong to another species group. Leucochrysa (N.) confusa appears to be closely related to
L. (N.) maculata, but L. (N.) maculata has heavy dark markings below the antennae on the frons which
forms a complete or interrupted band across the frons and the male gonarcus has a short vertical plate
dorso-medially.

The name “confusa” comes from the Latin confusio meaning mixture or disorder, and refers to
the difficulty in interpreting the male genitalia of this species.

HEAD. — Yellow with red markings. Vertex pale, without markings, except red spot behind each
antennal base. Gena with red stripe extending slightly onto frons. Frons pale, without markings. Scape
with dorso-lateral dark brown stripe; pedicel pale with apical dark ring; flagellum pale.

THORAX. — Cervical sclerite red (Fig. 61A,C). Pronotum green with thin lateral red stripe on an-
terior half. Mesonotum with irregular dark marks on prescutum and scutum. Metanotum pale, without
markings (Fig. 61A). Wing: Forewing pterostigma with well-developed dark spot basally. Venation
pale, except costal and R-Rs crossveins, origin of Rs and Ma, inner gradates, Psm-Pscu crossveins,
apex of forked and unforked marginal veins, and anal veins dark. Origin of Rs, inner gradates, and last
Psm-Pscu crossvein shaded. Forewing length (1.3 cm): width (0.41 cm) = ratio (3.17). Hindwing ve-
nation pale, except costal, R-Rs crossveins and inner gradates dark. Length (1.12 cm): width
(0.34 cm) = ratio (3.29) (Fig.61B).

MALE ABDOMEN. — Green with red markings on tergites 2, 6, and 7 (Fig. 61G); densely covered
by long setae on tergites and sternites (Fig. 61H). Microtholi absent. Gonarcus arched, thick with
short, narrow lateral arms. Gonocornua elongate, apically forcipate and embracing arcessus. Arcessus

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 297

slender and apically decurved with apical hook flanked by vertical plate-like lobes. Gonosaccus with
numerous tiny gonosetae with large bases (Fig. 61, D, E, F).
MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) cornuta de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Guaira, SP, 12-XII-97, Angelini, M. R.” (corn).

DIAGNOSIS. — This is one of a group of species best defined by elements of the male genitalia.
The gonocornua are long and relatively straight, and the arcessus has a pair of subapical dorsal horns
and sharply decurved apex. Other members of this group include: L. (N.) clepsydra, L. (N.) cruentata,
L. (N.) lenora Banks (1944), and L. (N.) nictheroyana (Navas, 1926). Of these species L. (N.)
cruentata and L. (N.) nictheroyana have a completely dark dorsal surface of the scape, complete lat-
eral pronotal stripe, dark palpi, and curved, forcipate gonocornua. Leucochrysa (N.) lenora is a more
northern species with red spots below each antenna on the frons, but unlike L. (N.) cornuta the
antennal scape dorsally is heavily marked with a dorso-lateral stripe that expands apically to encom-
pass the whole surface, the pronotum only has a pair of small dark spots at the anterio-lateral margins,
and the mesonotum has extensive dark markings along the anterior margin of the prescutum and later-
ally on the scutum. However, the male genitalia are almost identical. Even so, the non-genitalic char-
acter states are so different as to preclude these being regional variants.

The name “cornuta” comes from the Latin cornutus, which means horned, and refers to the
cone-like shape of the lateral arms of the gonarcus.

HEAD. — Green. Gena with dark red mark near frons (Fig. 62C). Frons pale, without markings.
Maxillary and labial palpi pale. Vertex pale, except for dark red continuation of scape stripe onto
antennal base. Scape with dorso-lateral dark red stripe; pedicel pale with apical dark ring; flagellum
pale (Fig. 62A).

THORAX. — Green. Pronotum pale with thin, lateral dark red stripe on anterior half. Mesonotum
pale with pair of small irregular dark marks on prescutum and scutum (Fig. 62A). Wings: Forewing
pterostigma with faintly dark basal spot. Venation green, except anterior junction of costal crossveins,
apical radial crossveins, middle part of Rs, inner and outer gradates, vertex of intramedian cell,
Psm-Pscu crossveins, and hind marginal crossveins dark (Fig. 62B). Length (1.54 cm): width
(0.56 cm) = ratio (2.75). Hindwing venation green, except costal and R-Rs crossveins dark (Fig. 62B).
Length (1.30 cm): width (0.43 cm) = ratio (3.02).

ABDOMEN. — Green; tergites with red markings; sternites 2—8 with numerous microtholi. Male:
Gonarcus strongly arched, with ovate lateral arms. Gonocornua elongate, evenly tapered, slightly sin-
uous at mid-length. Arcessus with two well-developed subapical horns and decurved, apico-medial
hook. Gonosaccus with few scattered setae (Fig. 62D, E, F).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) forciformis de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Bra-MT-Itiquira, P. E.Michelin, 20/1/97, SP41B, <, Freitas, S.” (rubber).

DIAGNOSIS. — This species is a member of a group of species with bifurcate apices to the
gonocornua. Members of the group also have pale palpi, a dark dorsal abdominal stripe and dark
markings at the lateral margins of the meso- and metascutellum. However, L. (N.) forciformis can be
distinguished from L. (N.) forcipata and L. (N.) furcata by the broad lateral pronotal stripe, by the dark
transverse band behind the antennal bases, and broadly contiguous bases of the gonocornua.

298 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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The name comes from the Latin forceps meaning pincer-like, and forma meaning form, referring
to the shape of the bifurcate apices of the gonocornua.

HEAD. — Yellow. Clypeus and frons with diffuse red coloration. Gena pale, unmarked. Vertex
with broad, transverse brown band behind antennal bases; pair of indistinct parallel red stripes on
raised medial area (Fig. 63D). Maxillary and labial palpi pale. Scape with diffuse reddish brown spot;
pedicel pale with apical dark ring; flagellum black.

THORAX. — Yellow. Pronotum pale with broad, dark, lateral stripe. Meso- and metanota with
dark lateral stripe and lateral margin of meso- and metascutellum (Fig. 63A). Wings: Forewing
pterostigma with large dark spot basally. Venation green, except apex of Rs and adjacent crossveins,
gradates, forked and unforked marginal veins, anal veins, and hind margin dark. Veins in anal area
more intensively darkened. Length (1.88 cm): width (0.62 cm) = ratio (3.03). Hindwing venation
green, except apex of Rs, gradates, apices of forked and unforked marginal veins, and hind margin
dark. Anal region more intensively darkened (Fig. 63B). Length (1.65 cm); width (0.50 cm) = ratio
(3.30).

ABDOMEN. — Microtholi present. Tergites dark, forming dark medial stripe (Fig. 63C). Male:
Gonarcus thick; strongly arched in broad V-shape; lateral arms broadly flattened. Gonocornua with
broad, contiguous bases and bifurcate, decurved hooks apically. Arcessus short, without subapical
horns or plate, apico-medial decurved hook with poorly developed lateral lobes. Gonosaccus with
gonosetae on conspicuous conical bases in two lateral pockets, most medial gonosetae longer and
most lateral gonosetae shorter; area beneath gonocornua with small sclerotized pits on membrane
(Fig. 63E, F). Female: Spermatheca short, with many coils. Bursa with pair of filamentous accessory
glands (Fig. 63G).

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 17 July 1995, Freitas, S. (1?
paratype); Mato Grosso: Itiquira, 20 January 1997, Freitas, S. (2° paratypes)(rubber).

Leucochrysa (Nodita) furcata de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 20-I-97, Scomparin, C. H., V.216, SP41D” (rubber).

DIAGNOSIS. — This species is amember of the group of species easily distinguished by the bifur-
cate gonocornua. Within the group, this species has a shorter arcessus, and shorter gonocornua with
apical teeth of unequal length. Additionally, it does not have the development of dark coloration on
meso- and metascutellum that the nominate species has. It can be separated from L. (N.) forciformis by
the lack of dark band behind the antennae, lack of dark lateral stripes on meso- and metanota, and
widely separated bases of the gonocornua of L. (N.) furcata. However, the most distinctive feature
(found in no other species) is the pair of sclerotized sacs at the apex of sternite 8 + 9 in males of of L.
(N.) furcata.

The name “furcata” is derived from the Latin furca, meaning fork-bearing, which refers to the bi-
furcate apex of the gonocornua.

HEAD. — Yellow. Clypeus and frons with diffuse red pigmentation. Gena pale, unmarked. Ver-
tex diffuse red with anterior pair of short, convergent, dark stripes (Fig. 64F). Maxillary and labial pal-
pi pale. Scape with broad reddish spot; pedicel pale with apical dark ring; flagellum dark.

THORAX. — Green. Pronotum green with reddish brown stripe laterally. Mesoscutum and
scutellum pale with small lateral red mark. Metascutellum pale with dark red lateral mark, remainder
of metanotum pale green (Fig. 64A). Wings: Forewing pterostigma with well-developed dark mark
basally. Venation pale, except costal crossveins, R-Rs crossveins, Rs-Psm crossveins, Psm-Psc
crossveins, apex of Rs, gradates, marginal forks and anal veins dark. Length (0.93 cm): width
(0.31 cm) = ratio (3.00). Six inner and eight outer gradate crossveins (Fig. 64B). Hindwing acute.
Pterostigma with well-developed dark mark basally. Venation pale, except R-Rs crossveins, gradates,

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) Ue ls)

marginal forks, apex of Rs, and hind marginal veins dark (Fig.64B). Length (0.81 cm): width
(0.25 cm) = ratio (3.24). Five inner and eight outer gradate crossveins.

ABDOMEN. — All tergites dark, forming longitudinal dark stripe (Fig. 64C). Male: Apex of
sternite 8 + 9 with pair of laterally raised, sclerotized sacs and tiny medial denticles (Fig. 64 D, E).
Gonarcus slightly arched, thick, with rounded lateral arms. Gonocornua short, with divided,
two-pronged apex. Arcessus narrow, elongate, without subapical horns or plate, with apical decurved,
medial hook, but lateral lobes not well developed. Gonosaccus with numerous gonosetae (Fig. 64G,
H). Female: Spermatheca short, once coiled. Bursa with tiny denticles and pair of filamentous acces-
sory glands (Fig. 641, J).

OTHER MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 18 August 1998, Freitas, S.
(12 paratype)(rubber).

Leucochrysa (Nodita) guataparensis de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Luiz Antonio, SP, Faz. Guatapara, T. 22, Celpav, 27/07/92, Freitas, S.” (eucalyptus)

DIAGNOSIS. — This species has a bird’s-head-shaped arcessus. The only two other species with
this shape are L. (N.) /ancala, from which it can be distinguished by the pale palpi and broad and api-
cally decurved gonocornua, and L. (N.) affinis. The oblique pronotal markings and flattened
gonocornua probably indicate a close relationship between these two species. They can be separated
by the dark flagellum and broadly triangular shape of the gonocornua of L. (N.) guataparensis.

The name “guataparensis” refers to the small town of Guatapara, in which the primary type speci-
men was collected.

HEAD. — Yellow. Gena red. Vertex pale with V-shaped red mark behind antennal bases. Frons
pale with small red spot between and below antennae. Maxillary and labial palpi pale. Scape pale with
two dorsal red stripes medially fused; pedicel pale with apical dark ring; flagellum pale, but basal
antennomeres darkly shaded on anterior surface (Fig. 65A, D).

THORAX. — Green. Pronotum pale with dark red lateral stripe; pair of thin, oblique dark lines
fused to lateral stripes at posterio-lateral corners. Mesonotum pale with red mark at lateral margin of
prescutum (Fig. 65A). Metanotum pale yellow, without markings. Wings: Forewing pterostigma
heavily pigmented basally. Venation green, except costal crossveins, middle part of Rs and adjacent
crossveins, inner and outer gradates, apices of unforked and first forked marginal veins dark (Fig.
65B). Length (1.58—1.78 cm); width (0.53—0.64 cm). Hindwing pterostigma darker basally than in
forewing. Venation green, except costal crossveins, middle part of Rs, inner and outer gradates and
forked marginal veins dark (Fig. 65B). Length (1.47 cm): width (0.50 cm) = ratio (2.94).

ABDOMEN. — Tergites pale, with red markings (Fig. 65C). Male: Microtholi present. Gonarcus
only slightly arched, lateral arms small, elongate triangular. Gonocornua, in lateral view, sharply el-
bowed and broadly expanded at mid-length, apically decurved; in dorsal view, a broad triangular plate
with apical point. Arcessus short, broadly triangular, with subapical bird’s-head-shaped curve, in lat-
eral view; apically decurved as small medial hook and undeveloped lateral lobes (Fig. 65E, F). Fe-
male: Subgenitale well-sclerotized, with pair of heavily sclerotized lateral tubercles caudally.
Spermathecae short, sinuous. Bursa with pair of filamentous accessory glands (Fig. 65G, H, I).

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Luiz Antonio, | June 1994, Freitas, S.(1¢
paratype)(eucalyptus); Jaboticabal, S. (1 2 paratype), 30 March 1996, Freitas, S. (1 2 paratype); 4 De-
cember 1996, Freitas, S. (12 paratype); Mato Grosso: Itiquira, 20 January 1997, Freitas, S. (1¢
paratype)(rubber); 17 February 1997, Scomparin, C. H. J. (4c paratypes)(rubber).

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Leucochrysa (Nodita) ictericus de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Birigui, SP, 21-I-96, Scomparin, C. H. J.” (corn).

DIAGNOSIS. — Leucochrysa (N.) ictericus is closely related to L. (N.) rodriguezi. Both species
have a basal tooth on the gonocornua, which is found in no other species. They can be separated by the
more dorsal stripe on the scape and larger, more circular spot on the mesoscutum of L. (N.) ictericus.
However, the most apparent differences are in the male genitalia, where L. (N.) ictericus has an un-
usual gonarcus with lateral arms that project laterally, not anteriorly, with no arch, and the presence of
a medio-dorsal lobe. Neither of these characteristics 1s found in any other species of Leuwcochrysa
(Nodita). The gonocornua are also distinctively elongate, with an apical notch.

The name comes from the Greek ikterikos meaning jaundiced or yellow and refers to the basic
body coloration.

HEAD. — Yellow with red markings. Vertex pale with pair of oblique crescent-shaped red marks
on raised medial area. Frons pale with small red spot between antennal bases. Gena red, extending as
red suffusion on clypeus. Maxillary and labial palpi amber. Scape with mid-dorsal red stripe, pedicel
pale with apical black ring; flagellum dark, with basal antennomeres intensely black ventro-laterally
(Fig. 66A, C, D).

THORAX. — Yellow. Legs pale. Pronotum pale with lateral red stripes extending to lateral tip of
mesoprescutum. Mesoscutum with submedial, circular red spot (Fig. 66A). Metanotum pale, without
markings. Wings: Forewing pterostigma with faint dark spot basally. Venation pale, except costal
crossveins, R-Rs crossveins, Rs-Psm and Rs-Ig crossveins, and gradates dark. Gradate series diver-
gent. Length (1.55—1.69 cm); width (0.5 cm). Hindwing pterostigmatic spot faint. Venation pale, ex-
cept middle costal crossveins, R-Rs transverse veins, middle part of Rs and posterior inner gradates
dark; darkness of crossveins not as intense as on forewing. Length (1.33—1.47 cm); width
(0.39-0.47 cm) (Fig. 66B).

ABDOMEN. — Yellow with faint red spots. Male: Gonarcus not arched; lateral arms flat, ovate, at
right angle to arcessus; dorso-medial, caudally-directed lobe with apical tooth. Gonocornua elongate,
tips bifid; large, apically-directed basal tooth. Arcessus broad, with decurved apical hook flanked by
apically truncate membranous lobes. Gonosaccus with sparse gonosetose (Fig. 66E, F, G). Female:
Spermatheca short with long spermathecal duct (Fig. 661). Subgenitale with posterio-lateral lobes and
long anterio-medial projection (Fig. 66J, H).

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Birigui, 21 November 1996, Scomparin,
C. H. J. (1% paratype)(corn); Taquaritinga, 4 September 1993, Xavier, A. L. Q.(1ol ? paratypes)(or-
ange); Jaboticabal, 13 February 1991, Freitas, S. (1 2 paratype)(CAS); 9 August 1996, Freitas, S.(1¢
paratype)(orange); March 1996, Freitas, S. (1%32 paratypes)(SDF/CAS); 30 June 1992, Pessoa, R.
(1o paratype); 27 October 1995, Freitas, S. (12 paratype); 23 March 1996, Fernandes, O. A. (1?
paratype); 18 March 1998, Freitas, S. (1 paratype); 14 March 1995, Seguim, L. D. (1? paratype);
Nova Europa, 21 October 1996, Bergman, L. G. (1? paratype)(rubber).

Leucochrysa (Nodita) incognita de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 30-XII-96, Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — The short, twisted shape of the gonocornua is distinctive. We have not seen this
form in any other species and this feature should be sufficient for recognition of the species.

The name “incognita” comes from the Latin incognitus meaning unknown or strange, and refers
to the bizarre shape of the male gonocornua.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 301

HEAD. — Yellow. Frons and vertex without markings. Gena red (Fig. 67C). Maxillary
palpimeres pale, except four and basal half of five dark. Scape faintly darkened; pedicel pale with api-
cal dark ring; flagellum dark.

THORAX. — Pale, unmarked. Wing: Forewing pterostigma with dark mark basally. Venation
green, except first 15 costal crossveins, base and apex Rs, and inner and outer gradates dark. Length
(1.47 cm): width (0.55 cm) = ratio (2.67). Hindwing pterostigma with well-developed, dark mark ba-
sally. Venation pale, except gradates and costal crossveins somewhat dark (Fig. 67A). Length
(1.19 cm): width (0.39 cm) = ratio (3.05).

ABDOMEN. — Pale, unmarked. Dorsal apodeme of ectoproct heavily sclerotized and forked near
callus cerci. Apical half of sternite 8 + 9 strongly tapered (Fig. 67B). Male: Lateral arms of gonarcus
ovate, with irregular margins; strongly arched and curved. Gonocornua short, twisted. Arcessus nar-
row with straight medial hook flanked by small lateral lobe. Gonosaccus with sparse gonosetae (Fig.
67D).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) interata de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Jaboticabal, SP, 20-X-93, Marucci, R. (cotton).

DIAGNOSIS. — The combination of character states which help define this species include: thin,
red marks from frons to tentorial pits, dark spots on mesonotum, and pronotum with medial spots or
lateral stripe extensions. Three other species also share this suite of character states: L. (N.) affinis, L.
(N.) camposi, and L. (N.) parallela. Unlike L. (N.) interata, L. (N.) affinis has dark dorsum of the
scape, dark antennae, and no dark metanotal markings. Leucochrysa (N.) parallela also has no dark
metanotal markings and no medial spots on the pronotum. Probably the most closely related species 1s
L. (N.) camposi, which has more extensive markings of meso- and metanotum and abdominal tergites
than L. (N.) interata. Additionally, the slender elongate spermatheca and broad antero-medial projec-
tion of the subgenitale are distinctive for L. (N.) interata.

The name “interata” comes from the Latin inter meaning between and atus meaning pertaining
to, and refers to the apex of the spermathecal gland which is neither hairy (the usual state for
chrysopids) nor bearing small plates.

HEAD. — Yellow. Frons and clypeus unmarked. Gena with small red spot near frons. Maxillary
and labial palpi pale. Vertex yellow with pair of crescent-shaped red stripes on raised medial area.
Scape and pedicel with red stripe dorso-laterally; flagellum pale, with basal antennomeres dark on lat-
eral margin (Fig. 68A, C, D).

THORAX. — Pronotum green, with pair of black punctures at mid-length; red stripe weakly de-
fined laterally. Mesoprescutum and scutum pale, with pair of diffuse red marks. Metanotum pale, with
pair of pale red marks (Fig. 68A). Wing: Pterostigma with faint dark mark basally. Forewing venation
green, except apical forks and basal radial and medial crossveins dark. Inner and outer gradates pale
(Fig. 68B). Forewing length (1.93 cm): width (0.72 cm) = ratio (2.68). Hindwing length (1.64 cm):
width (0.54 cm) = ratio (3.04).

ABDOMEN. — Pale; tergites | and 2 with red marks (Fig. 68E). Female: Spermatheca long; ven-
tral impression short. Surface of spermathecal duct apex smooth, neither hairy nor with tiny plates.
Subgenitale with broad, heavily-sclerotized antero-medial projection (Fig. 68F, G).

MATERIAL EXAMINED. — Known only from the holotype.

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Leucochrysa (Nodita) lineata de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Luiz Antonio, SP, Fz. Guatapara, T.25, Celpav, 0, 1-X-92, SP41, Freitas, S.” (eucalyptus).

DIAGNOSIS. — There is a group of species that has dark abdominal tergites that form a continu-
ous dark dorsal stripe and apically bifurcate gonocornua of the male genitalia. Among these species
there appears to be a progression from L. (N.) rodriguezi which has only crescent-shaped marks on the
abdomen, arched gonarcus, and narrow gonocornua with only a slight indication of apical bifurcation;
to L. (N.) ictericus with a complete abdominal stripe, flattened gonarcus with lateral arms at right an-
gles to the arcessus, and narrow gonocornua with slightly bifurcate apices; to L. (N.) lineata with
gonocornua greatly swollen basally and apically cleft (or hooked). Intermediate stages of this devel-
opment can be observed in the male genitalia of L. (N.) michelini, L. (N.) retusa, and L. (N.) tabacinus.
However, the differences among individual species in this grouping are so striking that they could not
be considered as variants of a single species. This group of species appears to be confined to southern
South America.

The name “lineata” is derived from the Latin /inea meaning line or thread, and refers to the dark
medial line on the abdominal tergum of this species.

HEAD. — Yellow. Maxillary and labial palpi amber. Gena pale, unmarked. Indistinct reddish
brown transverse band at clypeal-frontal suture; small dark red spot below antennae on mid-line. Ver-
tex with pair of divergent red stripes, which at anterior end form dark red double spot. Scape suffused
red dorsally, pale ventrally; pedicel pale with apical dark ring; flagellum black (Fig. 69C).

THORAX. — Pronotum green, with broad brown stripe laterally. Mesoprescutum brown laterally
along prescutal-scutal suture; scutum with two interconnected brown marks on either side; scutellum
with post-lateral brown dot. Metascutum and scutellum with large brown marks submedially (Fig.
69A). Wing: Forewing pterostigmal spot faint. Venation green, except origin of Rs, Rs vein for first
five cells and contiguous crossveins, gradates and marginal forks dark. Length (1.58 cm): width
(0.56 cm) = ratio (2.82). Hindwing venation pale, except apex of Rs and contiguous crossveins,
gradates and marginal forks dark (Fig. 69B). Length (1.41 cm): width (0.45 cm) = ratio (3.13).

ABDOMEN. — Tergites dark, forming dark mid-line (Fig. 69D). Microtholi present on sternite
8 + 9. Male: Tiny lanceolate gonocristae at apex of sternite 8 + 9. Gonarcus slender, only slightly
arched; lateral arms almost in straight line. Gonocornua basally greatly swollen, with medially
curved, apico-lateral hook. Arcessus with apically-decurved medial hook flanked by pair of rounded,
membranous lobes. Gonosaccus with gonosetae on conspicuous conical bases in pair of lateral pock-
ets; anterior to field of gonosetae, field of tiny linear pits below gonocornua on lateral margins of
gonosaccus (Fig. 69E, F).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) maculosa de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Taquaritinga, SP, Brasil, I-93, Fz. Sao Jose, Xavier, A. L. Q.” (orange).

DIAGNOsIs. — Almost all species of L. (Nodita) with heavy shading of forewing crossveins [e.g.,
L.(N.) postica] also have a dark apex of the hindwing. The heavy shading of forewing crossveins com-
bined with pale apex of the hindwing is unusual. Of the species with heavy shading of the forewing
crossveins, only L. (N.) maculosa has a flagellum which is basally dark and pale after the first two seg-
ments. The short spermatheca is also unknown in other species of Leucochrysa. In this respect, it more
closely resembles species of Chrysopini with cordate spermatheca, such as Chrysoperla externa and
Plesiochrysa brasiliensis.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 303

The name “maculosa,” from the Latin macula meaning spot or mark, refers to the heavily
margined crossveins in the forewing, which gives this species the appearance of having spotted
forewings.

HEAD. — Pale yellow. Frons green with wine red marks below antennal bases from eye margin to
medial margin of antennae and ventrally to below anterior tentorial pit. Vertex green with antennal
fossa wine red. Maxillary palpi pale basally, dark on fourth and basal half of apical segment, pale on
apical half (Fig. 70D). Scape with dorso-lateral and medial red stripes fused dorsally; pedicel pale
with apical dark ring; first two antennomeres dark, apical segments pale (Fig.70A).

THORAX. — Cervical sclerites red. Pronotum green with pair of oblique wine red marks on poste-
rior 2/3. Mesoprescutum pale with brick red mark along lateral margin, extending onto scutum near
wing base. Metanotum pale, without markings (Fig. 70A). Wings: Fore- and hindwing pterostigma
with well-developed dark mark basally. Venation green, except origin of Rs, apical radial crossveins,
Psm-Psc crossveins, gradates and apical forks dark and heavily shaded. Length (1.47 cm): width
(0.54 cm) = ratio (2.72). Hindwing venation green (Fig. 70B). Length (1.24 cm): width (0.40 cm) =
ratio (3.10). Tarsal claws without basal expansion.

ABDOMEN. — Green, with red spot on second tergite. Female: Spermatheca short, thick, not
coiled; ventral impression broad basally (Fig. 70C).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) michelini de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP, XI-96, Tosi, E. &”.

DIAGNOSIS. — This species is a member of a group having dark abdominal tergites and flattened
gonocornua with apical hook [see discussion under L. (N.) lineata]. Of the members of this group with
swollen bases of the gonocornua, L. (N.) tabacinus has less enlarged gonocornua with bifid apex (not
hooked), and broadly widened arch of the gonarcus. The marks on the vertex also appear smaller and
on metanotum much more developed in L. (N.) tabacinus. The most similar species to L. (N.)
michelini is L. (N.) lineata, which has the same inflated gonocornua, but the lateral arms of the
gonarcus have no arch, forming a straight line perpendicular to the arcessus. Leucochrysa (N.) lineata
also appears to have more dark pigmentation on the mesonotum than L. (N.) michelini.

The name was given in honor of P. E. Michelin who has given financial support to the senior au-
thor for chrysopid research.

HEAD. — Yellow. Vertex pale with U-shaped red mark on raised medial area interrupted at
mid-line and darker anteriorly. Scape with diffuse red spot dorsally; pedicel pale with apical dark ring;
flagellum black. Maxillary and labial palpi amber. Gena pale, unmarked. A diffuse red transverse
band on either side of clypeal-frontal suture. Small red spot below and between antennae (Fig. 71D).

THORAX. — Pronotum green with broad brown lateral stripe. Middle part prescutum dark; two
crescent-shaped marks on mesoscutum. Submedial part of metascutum and lateral portion of
scutellum dark brown (Fig. 71A). Wing: Forewing pterostigma slightly darkened basally. Venation
green; except most of costal crossveins, apex of Rs and contiguous veins, gradates and marginal forks
dark. Length (1.68 cm): width (0.58 cm) = ratio (2.90). Hindwing apically acute. Pterostigma with
faint darkening basally. Venation green, except apex of Rs and contiguous veins black (Fig. 71B).
Length (1.44 cm): width (0.45 cm) = ratio (3.20).

ABDOMEN. — Microtholi present. Tergites dark, forming dark mid-line (Fig. 71C). Male:
Gonarcus strongly arched with rounded lateral arms. Gonocornua broadly expanded plates with api-
cal decurved hook. Arcessus short; subapical horns and plate absent; decurved apico-medial hook
present with lateral lobes flattened or absent. Gonosaccus with numerous setae on large conical bases

304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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arranged in two lateral fields; field of tiny linear marks present near gonocornua, and directly below
gonocornua field of tiny punctures (Fig. 71E, F).

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, 26 October 1995, Ribeiro,
M. C. (1c paratype)(orange).

Leucochrysa (Nodita) parallela de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Bra-SP-Jaboticabal, FCAV, 10-ITI-99, SP58, o’, Freitas, S.” (cotton).

DIAGNOSIS. — The lateral arms of the gonarcus of this species are flattened, rather than vertical,
as in other species. The lack of extensive dark markings on the thorax, especially the lateral portion of
the metascutellum and minimal abdominal tergite markings also suggest a distant relationship with
the forcipata group. We know of no other species with such strikingly flattened lateral arms.

The name “parallela” comes from the Latin parallelus meaning side by side equidistantly, and re-
fers to the parallel, flattened arms of the gonarcus.

HEAD. — Yellow. Frons and clypeus pale, unmarked. Gena pale with small red spot along frontal
suture. Maxillary and labial palpi pale (Fig. 72A, D). Vertex pale, unmarked. Antennal scape pale,
with red lateral stripe thin at midlength; flagellum pale.

THORAX. — Pronotum green with lateral red stripe thinner at mid-length. Mesoscutum pale with
pair of circular red spots; second axillary sclerite with small red spot (Fig.72A). Wings: Forewing
pterostigma faintly darkened basally. Venation green, except costal ends of costal crossveins, middle
of R- Rs crossveins, first five Rs-Psm crossveins, gradates, Psm-Psc crossveins, forked and unforked
marginal veins, and apices of anal veins dark (Fig. 72B). Length (1.40 cm): width (0.50 cm) = ratio
(2.80). Hindwing pterostigma heavily pigmented basally. Venation green, except middle part of Rs
and gradates dark (Fig. 72B). Length (1.19 cm): width (0.40 cm) = ratio (2.98).

ABDOMEN. — Tergites pale, with red spots on tergites 2 and 3. Male: Gonarcus short, with lateral
arms large, rounded, truncate posteriorly, with latero-apical, decurved point. Arcessus angulate in lat-
eral view; distal half with thin, semimembranous and apically divergent ridges; terminal hook flanked
by paired membranous lobes (Fig.72E, F). Ectoproct heavily sclerotized; ventral branch of dorsal
apodeme extends to postventral corner of ectoproct as highly sclerotized, obtuse point. Microtholi ab-
sent.

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Jaboticabal, October 1992, Delfino, T. H.
(1c paratype)

Leucochrysa (Nodita) retusa de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
“Balsamo, SP, 29-IX- September 1997, Bergman, E.” (rubber).

DIAGNOSIS. — Males of this species appear to have a less developed state of swollen gonocornua
and hooked or bifurcate apex, such as found in L. (N.) lineata and L. (N.) tabacinus. Perhaps the most
similar species is L. (N.) furcata, which is part of the group of species with bifid gonocornua. Like L.
(N.) retusa, L. (N.) furcata has short, relatively broad gonocornua, and relatively few meso- and
metanotal markings. However, L. (N.) retusa does not have the darkened abdominal tergites of this
species. The thin, quadrate-shaped gonarcus with small lateral arms and broad arcessus are distinctive
for L. (N.) retusa.

The name “retusa” comes from the Latin retusus meaning blunted or notched, and refers to the
unusual squared arch of the gonarcus.

a

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HEAD. — Yellow, without marks. Maxillary palpimeres basally pale; third, fourth and half of
fifth segments dark. Scape pale with dark dorso-lateral mark; pedicel pale with dorso-lateral dark
mark; flagellum pale (Fig. 73A).

THORAX. — Cervical sclerite red. Pronotum with thin dark red stripe laterally on anterior 2/3.
Meso- and metanotum pale, unmarked (Fig. 73A). Wings: Forewing pterostigma faintly darkened ba-
sally. Venation green, except junctures of costal crossveins, R-Rs crossveins, gradates, and marginal
forked veins dark (Fig. 73B). Length (1.44 cm): width (0.53 cm) = ratio (2.72). Hindwing pterostigma
dark basally. Venation pale, except middle part of Rs and gradates dark. Length (1.24 cm): width
(0.41 cm) = ratio (3.02).

ABDOMEN. — Pale, except tergite 2 with pair of triangular red spots. Male: Medial arch of
gonarcus thin, quadrate. Gonocornua widely spaced, flattened, with apico-lateral hook. Arcessus
broad, with subapical transverse ridge; decurved apico-medial hook with poorly developed lateral
lobes. Gonosaccus with few, scattered, large gonosetae (Fig. 73C, D).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) robusta de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, SP, 7-IX-99, PEM, 28, Freitas, S.” (rubber).

DIAGNOSIS. — This species shares with L. (N.) santini a dorsal scape stripe, dark pronotal stripe
which expands posteriorly, absence of meso- and metanotal markings, and quite short gonocornua.
However, L. (N.) santini has dark basal palpimeres and almost straight gonocornua in dorsal view,
which are also slightly larger and more closely spaced than in L. (N.) robusta.

The name “robusta” comes from the Latin word robustus, meaning hard and strong like an oak,
and refers to the thick arch of the gonarcus with broad arcessus.

HEAD. — Pale yellow. Gena pale, with transverse red band close to frons. Frons green, without
markings. Maxillary palpimeres pale (Fig. 74A). Vertex green, without marks (Fig. 74A, D). Scape
pale, with dorso-latersal red stripe not reaching dorsal and ventral margins; pedicel pale, with dark
marks; flagellum pale.

THORAX. — Green. Pronotum pale, glabrous, with irregular red mark on lateral margin expanded
posteriorly (Fig. 74A). Mesonotum and metanotum pale, without markings. Wings: Forewing
pterostigma faintly darkened basally. Venation green, except crossvein endings, and inner and outer
gradates dark. Apex of intramedian vein reaches Psm after first Rs-Psm crossvein. Length
(1.33—1.51 cm); width (0.5—0.56 cm). Hindwing pterostigma darkened basally. Venation green, ex-
cept costal crossveins, radial crossveins 7-9, apex of Psm, and inner and outer gradates dark. Length
(1.17—1.25 cm); width (0.30—-0.43 cm) (Fig. 74B).

ABDOMEN. — Green. Tergites with dark red marks. Sternites 2-8 with microtholi. Male:
Gonarcus large, thick, strongly-arched, with ovate lateral arms. Gonocornua small, laterally curved.
Arcessus broad; without subapical horns or plate; decurved apico-medial hook with lateral lobes
forming two small ridges (Fig.74F). Female: Spermatheca cordate at apex, continuing to mass of con-
voluted tubes; spermathecal duct contorted, terminally expanded; ventral impression wide, swollen;
accessory glands short, stout, tubular. Subgenitale short; with thin-walled, pocket-like medial exten-
sion (Fig. 74G, H).

OTHER MATERIAL EXAMINED. — Brazil: S40 Paulo: Jaboticabal, 19 November 1995, Freitas,
S. (12 paratype); Luiz Antonio, 13 August 1992, Freitas, S. (1% paratype)(corn); 27 July 1992,
Freitas, S. (1@%22 paratypes); Mato Grosso: Itiquira, 7 September 1999, Freitas, S. (1%
paratype)(rubber).

306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Leucochrysa (Nodita) santini de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Jaboticabal, SP, 28-VII-1995, Freitas, S.”

DIAGNOSIS. — This species has few distinctive characteristics. The shape of the lateral pronotal
stripe, similar abdominal markings and small unmodified gonocornua would indicate a closer rela-
tionship with L. (N.) robusta than with other species. The two species can be separated by the dark pal-
pi and straight shape of the gonocornua of L. (N.) santini.

The name was given in honor of Prof. Santin Gravena who provided some specimens for this
study.

HEAD. — Pale yellow. Gena red. Frons green, without dark markings. Maxillary palpimeres pale
basally; palpimeres 3-4 and basal part of 5 dark; apex of palpimere 5 pale. Vertex green, without
markings (Fig. 75A, D). Scape with dorso-lateral red stripe, not reaching basal or apical margins (Fig.
75A); pedicel pale with dark marks; flagellum pale.

THORAX. — Green. Pronotum pale with bright red marginal stripe abruptly broadened posterior
to mid-length (Fig. 75A). Mesonotum pale with dark red markings. Metanotum pale, without mark-
ings. Wings: Forewing pterostigma faintly darkened basally. Venation green, except junctions of
crossveins, and inner and outer gradates dark. Apex of intramedian cell reaches Psm after the first
Rs-Psm crossvein. Length (1.28—1.58 cm); width (0.47—0.58 cm) (Fig. 75B). Hindwing pterostigma
strongly darkened basally. Venation green, except costal crossveins and gradates dark (Fig.75B).
Length (1.21 cm): width (0.40 cm); ratio = (3.03).

ABDOMEN. — Green. Tergites with dark red markings on segments 3, 4, 5, 7, and 8 (Fig. 75C).
Sternites 2-8 with many microtholi. Male: Gonarcus thick, strongly arched, with vertically oriented,
ovate lateral arms. Gonocornua small, straight, apically decurved. Arcessus without subapical horns
or plate; apically decurved hook flanked by small, lateral ridges. Gonosaccus with scattered
gonosetae (Fig. 74E, F). Female: Spermatheca multiply-coiled, ventral impression small with many
turns (Fig. 34G).

OTHER MATERIAL EXAMINED. — Brazil: Sao Paulo: Jaboticabal, August 1996, Freitas, S. (1$
paratype)(CAS); Luiz Antonio, 25 August 1992, Freitas, S. (2% paratypes)(eucalyptus); 27 July 1992,
Freitas, S., (1% paratype)(eucalyptus)(CAS); Minas Gerais: Belo Horizonte, 2 September 1991,
Kumagai, A. (2% paratypes).

Leucochrysa (Nodita) scomparini de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Itiquira, MT, 18-XI-96, Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — Females of this species are immediately recognizable and separable from all
other known species by the apical, tight double coil, which forms a knob at the end of the spermatheca
(Fig. 76E). The extensive dark meso- and metanotal markings and dark abdominal tergites relate this
species to a group in which males have enlarged, flattened gonocornua that are apically bifid or
hooked [see discussion of L. (N.) lineata].

This species is dedicated to the enthusiastic entomologist Cassio Henrique Junqueira Scomparin,
who has attempted to use chrysopids in biological control of agricultural pests.

HEAD. — Yellow with red markings. Vertex pale, with lateral red stripe close to ocular margins;
convergent crescentic stripes on raised medial area (Fig. 76A). Scape pale, with red dorso-lateral
stripe and short medial stripe; pedicel pale with red ring; flagellum pale. Gena red. Frons pale with
small red spot below and between antennal bases. Maxillary and labial palpi pale (Fig. 76C).

THORAX. — Pronotum yellowish green, with red spots at the anterio-lateral corner and pair of red
spots at posterio-medial margin. Mesonotum pale with extensive red and brown marks, prescutum

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 307

medially with red spots and laterally brown; scutum and postscutum with reticulate brown marks;
mesoscutellum pale medially, brown laterally. Metanotum broadly brown (Fig. 76A). Wings:
Forewing and hindwing pterostigma with dark brown spot basally. Venation pale, except costal junc-
ture of costal crossveins, origin of radial sector, middle part of Rs, gradates, forked marginal veins,
and apices of marginal unforked veins dark. Length (1.66 cm): width (0.66 cm) = ratio (2.52).
Hindwing venation pale, except costal crossveins, middle part of Rs, gradates and forked marginal
veins dark (Fig. 76B). Length (1.44 cm): width (0.48 cm) = ratio (3.00).

ABDOMEN. — Green. Tergites 4~7 with large, dark spots (Fig. 76A). Female: Spermatheca
short, vela with two tight apical rings. Bursa much folded, with pair of filamentous accessory glands.
Subgenitale wide, with short anterio-medial projection (Fig. 76D, E).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) squamisetosa de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
“Birigui, SP, Fz. Sao Joaquim, | 1/June/94, SP68, Scomparin, C. H. J.” (corn).

DIAGNOSIS. — The swollen and darkened, second anal vein of the forewing is similar to the basal
markings of Ceraeochrysa tauberae from Costa Rica (Penny 1997), but is otherwise unknown in L.
(Nodita). The contorted subapical ridges of the arcessus is also a characteristic unique to this species.
The extensive red markings of the head are not often seen in this subgenus, but there are a few other
species with similar markings, such as L. (N.) aleura (Banks, 1944), L. (N.) morrisoni (Navas, 1914),
and L. (N.) trifurcata (Banks, 1948). However, no other species of L. (Nodita) has the distinctive,
thickened gonosetae, some of which have basal plates.

The name “squamisetosa” comes from the Latin sqguama and seta meaning scaly hairs, and refers
to the scattered large gonosetae on large conical bases or flat plates of the gonosaccus.

HEAD. — Pale yellow. Clypeus pale, marked with narrow red band between eyes through
clypeus under anterior tentorial pit; broader medially. Frons pale with narrow, red, double concentric
ring below antennal sockets fused at mid-line and continued dorsally as a narrow median stripe. Ver-
tex pale, with red partial ring at base of each scape continued postero-laterally as a narrow stripe along
eye margin; pair of submedial, parallel, red stripes. Labial and maxillary palpi pale. Scape with dorsal
surface red, ventral surface pale; pedicel and flagellum pale (Fig. 77A, C).

THORAX. — Pronotum wider than long, yellow-green, unmarked. Meso and metanotum pale
with diffuse reddish markings; no pale median stripe (Fig. 77A). Wings: Fore- and hindwings with
dark basal spot. Venation green, except crossveins and bases of longitudinal veins dark; no bordering.
Basal fork of second anal veins swollen, darkened and bordered. Forewing with five inner, and seven
outer gradates, the series converging posteriorly (Fig. 77B). Forewing length (1.20 cm): width
(0.50 cm) = ratio (2.40). Hindwing length (1.08 cm): width (0.34 cm) = ratio (3.18).

ABDOMEN. — Pale. Tergites 3, 4, 6 and 7 with broad dark marks (Fig. 77D). Apex of sternite 9
truncate. Dorsal apodeme of ectoproct extended dorso-medially at apex. Callus cerci black posteri-
orly. Male: Microtholi large, dense on sternites 3—8. Gonarcus thick, broadly arcuate; lateral arms tri-
angular. Gonocornua short, straight, apically obtuse and decurved. Arcessus broad basally, little
sclerotized laterally; with low, contorted dorsal ridges at subapical angle, apical hook small, with lat-
eral lobes transverse. Gonosaccus with gonosetae borne subapically on sclerotized, conical projec-
tions; ventral pair of thick gonosetae on oval scleritized bases (Fig. 77E, F, G).

MATERIAL EXAMINED. — Known only from the holotype.

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Leucochrysa (Nodita) tabacinus de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, 20-I-97, Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — This species is a member of the group of species with completely darkened ab-
dominal tergites and flattened, apically bifid gonocornua. The gonocornua, while well developed in L.
(N.) tabacinus, are not as swollen as in some other species in the group, such as L. (N.) lineata and L.
(N.) michelini. The lateral arms of the gonarcus, although broadly angled with respect to the medial
arch in L. (N.) tabacinus, do not form a straight line, as is found in L. (N.) ictericus and L. (N.) lineata.
The extensive brown coloration of thorax and abdomen is also seen in L. (N.) scomparini, but unlike L.
(N.) tabacinus, that species has red gena, completely dark metascutellum, and a distinctive dou-
bly-coiled knob on the spermatheca.

The name “‘tabacinus” comes from the New Latin for tabacum for tobacco and inus meaning per-
taining to, referring to the extensive tobacco brown coloration of the body.

HEAD. — Yellow. Clypeus and frons without markings, except a small dark spot between and be-
low antennal bases. Gena pale, unmarked. Vertex with short dark stripes antero-medially near
antennal bases. Antennal fossa dorsally red. Maxillary and labial palpi pale (Fig. 78C). Scapes pale
with diffuse reddish brown spot; pedicel pale, with apical dark ring; flagellum dark.

THORAX. — Pronotum yellow with lateral brown stripe not extended to anterior or posterior mar-
gins (Fig. 78A). Mesoprescutum pale with red mark antero-laterally; scutum pale with brown mark
posteriorly; mesoscutellum pale with lateral dark suffusion. Metascutum pale with pair of broad
submedial brown marks; metascutellum pale medially, with progressively darker suffusion laterally
(Fig. 78A). Wings: Fore- and hindwing faintly darkened basally. Venation green, except posterior
part of costal margin, outer gradates, marginal forks and unforked marginal veins, and anal veins dark;
posterior juncture of anal veins and hind margin intensely black. Length (3.06 cm): width (1.06 cm) =
ratio (2.89). Hindwing venation green, except apex of Rs and contiguous veins, and apical part of fore-
and hind margins dark; marginal junctures of anal veins and base of hind margin dark (Fig.78B).
Length (2.66 cm): width (0.84 cm) = ratio (3.17).

ABDOMEN. — Pale green, except all tergites with dark medial sclerites (Fig.78G). Male:
Gonarcus thin, curved, lateral arms continuation of same plane as medial arch. Gonocornua flattened,
with apex bifid, decurved. Arcessus short; subapical horns and plate absent; apical decurved medial
hook, flanked by rounded, enlarged lateral lobes. Gonosaccus with scattered gonosetae (Fig. 78E, F).
Female: Spermatheca short. Bursa convoluted and two thick accessory glands (Fig. 781).
Subgenitale highly sclerotized, with long antero-medial projection above large, round sclerotized
lobe (Fig. 78D, J)

OTHER MATERIAL EXAMINED. — Brazil: Mato Grosso: Itiquira, 20 June 1997, Scomparin,
C.H. J.” (20°22 paratypes)(rubber).

Leucochrysa (Nodita) tenuis de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Luiz Antonio, SP, Faz. Guatapara, T.25, Celpav, 6/12/93, Freitas, S.” (eucalyptus).

DIAGNOSIS. — This is a relatively pale species with few distinctive markings. However, the
gonarcus is quite distinct. No other species has the exaggerated, long, parallel lateral sides of the me-
dial arch of this species. The almost complete absence of gonocornua is also quite distinctive.

The name “tenuis” comes from the Latin meaning thin, and refers to incredibly long, thin, lateral
sides of the medial arch of the gonarcus.

HEAD. — Yellow. Gena pale, with diffuse red spot near frons (Fig. 79C). Frons pale, with small
red spot below and between antennal bases. Maxillary palpimeres basally pale; fourth and base of fifth

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 309

segments dark; apex pale. Vertex pale, unmarked (Fig. 79C). Scape pale, with dark brown
dorso-lateral stripe; pedicel and flagellum pale.

THORAX. — Green. Pronotum with dorso-lateral dark red stripe (Fig. 79A). Meso- and
metanotum pale, unmarked. Wings: Forewing pterostigma faintly darkened basally. Venation green,
except costal crossveins 7—13, apex of Rs, posterior apices of radial crossveins, and forked marginal
veins dark (Fig. 79B). Length (1.51 cm): width (0.56 cm) = ratio (2.70). Hindwing pterostigma with
well-developed dark spot basally. Venation green, except apex of Rs and forked marginal veins dark.
Length (1.34 cm): width (0.41 cm) = ratio (3.27).

ABDOMEN. — Green, without markings. Male: Sternites 2—8 with many microtholi. Gonarcus
medial arch quadrate, extremely long, thin, and parallel; lateral arms ovate. Gonocornua reduced to
small, rounded, basal lobes. Arcessus large, broad; subapical horns and plate absent; apical hook
well-sclerotized, not decurved, with lateral lobes well-developed, rounded. (Fig. 79D, E).

MATERIAL EXAMINED. — Known only from the holotype.

Leucochrysa (Nodita) vignisi de Freitas and Penny, new species

TYPE. — Male holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil, la-
beled “Itiquira, MT, 10-V-96, Scomparin, C. H. J.” (rubber).

DIAGNOSIS. — There are few distinctive features to this species. It appears to be part of a group of
species with poorly developed gonocornua and broad, bulging bases to the arcessus. The two species
most similar to L. (N.) vignisi appear to be L. (N.) retusa and L. (N.) santini, which also have thin, in-
complete stripes on the scape; pale vertex; pale meso- and metanotum; somewhat thin, incomplete lat-
eral pronotal stripes; and sparse, scattered gonosetae. Of the three, L. (N.) santini has dark palpi and
gena. The gonarcus of L. (N.) retusa is much thinner and more quadrate-shaped than in L. (N.) vignisi.
The shape of the gonocornua separates the three species most easily. The gonocornua of L. (N.) retusa
are relatively broad, with an apical hook along the lateral edge. The bases of the gonocornua of L. (N.)
santini are spaced much closer together than in L. (N.) vignisi.

This species was dedicated to Berthrand Vignis, who along with L. Barré enthusiastically opened
the way for chrysopid use for biological control of rubber plant pests.

HEAD. — Yellow. Frons, palpi, gena and vertex pale, unmarked. Scape pale, with incomplete,
thin dorso-lateral black stripe; pedicel and flagellum pale (Fig. 80A).

THORAX. — Pronotum green with medial yellow stripe and thin, lateral red stripe. Meso- and
metanotum pale, unmarked (Fig. 80A, C). Wings: Fore- and hindwing pterostigma with well-devel-
oped dark spot basally. Venation green, except costal junctures of costal crossveins, radial crossveins,
apex of Rs, anterior juncture of last medial crossvein, inner and outer gradates, Psm-Psc crossveins,
and forked and unforked marginal veins dark. Length (1.48 cm): width (0.56 cm) = ratio (2.64).
Hindwing venation green, except costal crossveins, radial crossveins, apex of Rs and forked marginal
veins dark (Fig. 80B). Length (1.30 cm): width (0.44 cm) = ratio (2.95).

ABDOMEN. — Green, with yellow mid-dorsal line. Tergites 3, 4, and 7 with dark spots (Fig.
80D). Male: Gonarcus strongly arched medially; lateral arms ovate. Gonocornua small, widely
spaced, apically forcipate. Entoprocessus broad, well developed; without subapically horns or plate;
apico-medial, decurved hook flanked by well-developed rounded ridges. Gonosaccus with many long
gonosetae (Fig 80E, F).

OTHER MATERIAL EXAMINED. — Brazil: Mato Grosso, Itiquira, 23 March 1997, Scomparin,
C. H. J.” (1% paratype)(rubber).

310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Leucochrysa (Nodita) vittatus de Freitas and Penny, new species

TYPE. — Female holotype, deposited at Museu de Zoologia/USP (MZUSP), Sao Paulo, Brazil,
labeled “Rib. Preto, SP, 2/11/91, Vieira, D. A.” (orange).

DIAGNOSIS. — This species has a rather atypical appearance for this subgenus. Usually, any dark
markings of the pronotum are in the form of lateral stripes and spots, sometimes coupled with oblique
stripes. Submedial pronotal stripes seem to be unique to this species. In many ways L. (N.) vittata re-
sembles L. (N.) scomparini with more extended and exaggerated submedial stripes. Both species have
extensive, similar dark markings of head and mesonotum. More extensive darkening of the abdominal
stripes found in L. (N.) vittata would create the condition seen in L. (N.) scomparini. However, there
are some significant differences between the two species. The metascutellum on L. (N.) vittata is pale,
while that of L. (N.) scomparini is dark. The dark scape stripe is dorsolateral in L. (N.) scomparini and
dorsomedial in L. (N.) vittatus while L. (N.) scomparini has an additional medial spot. The most dis-
tinct differences are in the female genitalia, where L. (N.) scomparini has a spermatheca with
tightly-coiled knob and non-glandular duct, whereas L. (N.) vittata has a larger, unknobbed
spermatheca and glandular spermathecal duct. Leucochrysa (N.) vittata also has an inner gradate se-
ries of the hindwing which runs close to the radial sector, while that of other species is more equidis-
tant between Rs and the outer gradate veins.

The name “vittatus” comes from the Latin vifta for ribbon, band or stripe and atus for provided
with, referring to the unusual number of brown stripes on the head and thorax.

HEAD. — Pale green. Gena brown. Frons pale, without markings. Maxillary and labial palpi pale
(Fig. 80A, D). Vertex pale with pair of dark parallel stripes on raised medial area, tapered
antero-medially to point between antennal bases; lateral pair of dark stripes along eye margin to poste-
rior margin of vertex. Scape with dorso-medial brown stripe continued briefly onto fossa; pedicel and
flagellum pale.

THORAX. — Pronotum with submedial brown stripes and small lateral spot each side halfway to
posterior margin. Mesoprescutum with three dark marks on either side, continued onto mesoscutum
and mesoscutellum, which have additional lateral stripes. Metanotum pale with submedial pair of
dark spots and postero-lateral oblique dark stripe. Pleura and metacoxae brown; metafemur pale
fuscous, brown banded apically (Fig. 80A). Wings: Pterostigma of both wings dark basally. Venation
green, except junctures of first ten costal crossveins and all of distal crossveins, radial crossveins dis-
tal to stigma, longitudinal veins at juncture with crossveins, and three cells of Rs at mid-length dark.
Forewing gradates parallel. Hindwing venation marked as in forewing. Inner series of gradates closer
to Rs than in forewing (Fig. 80B). Forewing length (1.59 cm): width (0.53 cm) = ratio (3.00).
Hindwing length (1.43 cm): width (0.43 cm) = ratio (3.33).

ABDOMEN. — Pale green. Tergites 2—8 with pair of submedial brown stripes (Fig. 80C). Female:
Spermatheca short, wide, with constriction between upper and lower chamber; vela region
lightly-sclerotized, moderately short; ventral impression short. Spermathecal duct wide, gently undu-
lated, glandular-walled region; longer than spermatheca. Bursa small. Bursal accessory glands not
seen. Subgenitale broadly rounded, dorsal lobes deeply incised medially; postero-medial lobe with
broad indentation and transverse lip (Fig. 80E, F, G).

MATERIAL EXAMINED. — Known only from the holotype.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE)

FIGURES

A list of abbreviations used with the figures is as follows:

acu = acumen

g = accessory glands

al = apical lobe

arc = arcessus

atp = anterior tentorial pit

b = copulatory bursa

bd = bursal duct

cc = callus cerci

cru = crumena

dapo = dorsal apodeme

dh = dorsal hood of the gonarcus
ent = entoprocessus

gc = gonarcus (medial arch)
gcn = gonocornua

gcr = gonocristae

gps = gonapsis

gsc = gonosaccus

gst = gonosetae

ig = inner gradates

imc = intramedian cell

la = lateral arm of the gonarcus
lg = lateral gonapophyses
m1 = first median cell

m2 = second median cell

mm = millimeters

mp = maxillary palpi

u = mediuncus

og = outer gradates

pa = paramere

Psc = pseudocubitus vein

Psm = pseudomedia vein

psp = pseudopenis

Rs = radial sector

S6 = sixth sternite

S7 = seventh sternite

S8 + 9 = fused eighth and ninth sternite
sap = subapical projection

sd = spermathecal duct

T7 = seventh tergite

T8 = eighth tergite

T9 + ect = ninth tergite and ectoproct
v = vela

vapo = ventral apodeme

vb = ventral branch of the dorsal apodeme
vi = ventral impression

311]

312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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LLL

FIGURE 1. Nacarina panchlora (Gerstaecker, | 888). A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D.
Maxillary palpi; E. Female genitalia; F. Apex of abdomen; G. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 313

A “ida © ff

0.5 mm

FIGURE 2. Nacarina pletorica (Navas, 1919). A. Head and thorax: dorsal view; B. Wings; C. Maxillary palpi; D. Head, fron-
tal view; E. Apex of male abdomen; F, G. Male genitalia, dorsal and lateral view.

314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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~\

IS
Wi

0.5mm

neri (Navas, 1924). A. Maxillary and labial palpi; B. Wings; C. Sternite 7; D. Subgenitalia; E.

FIGURE 3. Nacarina wag.
Spermatheca; F. Colleterial gland.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) SS

4
.

Ad ‘
voy
yete

ait

ww syeyt.

je)
ae
[e)

~

ars
v vee

spiny lobe 2
denticles +

ra iia

FiGure 4. Nacarina aculeata, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D. Apex of
male abdomen; E. Part of abdominal apex showing the denticles; F, G, H. Male genitalia, ventral, lateral and dorsal view, re-
spectively.

316 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 19

zea

|
FIGURE 5. Nacarina gladius, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D. Apex of
male abdomen; E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) a7

[LEZ

Ti

SU ueeereecwes<
Ta
wR Ss

FIGURE 6. Nacarina lavrasana, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D.
Maxillary and labial palpi; E. Apex of female abdomen; F. Subgenitale; G. Female genitalia; H. Male genitalia, dorsal view; I.
Male genitalia, lateral view.

318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 7. Nacarina sagitta, new species. A. Apex of male abdomen; B. Wings; C. Male genitalia, dorsal view; D. Male geni-
talia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 319

FIGURE 8. Ceraeochrysa acmon Penny, 1998. A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdomen; D.
Gonapsis; E. Male genitalia, lateral view; F. Male genitalia caudal view; G. Female genitalia; H. Subgenitale.

320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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T9+ect
dapo

al til a all —_err

0.5 mm

FIGURE 9. Ceraeochrysa caligata (Banks, 1945). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdomen;
D. Female genitalia; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Gonapsis; H. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 321

0.5mm

FIGURE 10. Ceraeochrysa cincta (Schneider, 1851). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdo-
men, lateral view; D. Apex of female abdomen; E. Female genitalia; F. Gonapsis; G. Subgenitale; H. Male genitalia, lateral
view; I. Male genitalia, dorsal view; da= dorsal apodeme of ectoproct, ent= entoprocessus, gps= gonapsis, vb= ventral branch
of dorsal apodeme.

322 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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\ a
).\'

GN cons
Sir

FIGURE 11. Ceraeochrysa claveri (Navas, 1911). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdomen;
D. Apex of female abdomen; E. Sternite 8 + 9, ventral view; F. Gonapsis; G. Female genitalia; H. Male genitalia, lateral view; I.
Male genitalia, dorsal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 323

gps 0.5 mm

FIGURE 12. Ceraeochrysa cubana (Hagen, 1861). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdomen,
lateral view; D. Apex of male abdomen, ventral view; E. Female genitalia; F. Male genitalia, lateral view; G. Male genitalia,
dorsal view.

324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 13. Ceraeochrysa everes (Banks, 1920). A. Head and prothorax, dorsal view; B. Wings; C. Male genitalia, lateral
view; D. Male genitalia, dorsal view; E. Detail of apex of arcessus; F. Subgenitale; G. Spermatheca; H. Gonapsis; I. Apex of ab-
domen.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 325

0.5 mm

FIGURE 14. Ceraeochrysa montoyana (Navas, 1913). A. Head and prothorax, dorsal view; B. Wings; C. Spermatheca; D.
Subgenitale.

326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Tn
L--Tq {il

0.5 mm

FIGURE 15. Ceraeochrysa paraguaria (Navas, 1919). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdo-
men, ventral view; D. Apex of male abdomen, lateral view; E. Gonapsis; F. Subgenitale; G. Female genitalia; H. Male genitalia,

lateral view; I. Male genitalia, dorsal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 327

LLL

2a NBER RS
>

1.0 mm
T9+ect

0.5 mm

FIGURE 16. Ceraeochrysa sanchezi (Navas, 1924). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdomen,
lateral view; D. Sternite 8 + 9; E. Subgenitale; F. Spermatheca, G. Gonapsis; H. Female genitalia; I. Male genitalia, lateral
view; J. Male genitalia, dorsal view.

328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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(7LUY

AS <4

FIGURE 17. Ceraeochrysa scapularis (Navas, 1914). A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Apex of male
abdomen, lateral view; D. Sternite 8 + 9; E. Gonapsis; F. Female genitalia; G. Subgenitale; H. Male genitalia, lateral view; I.
Male genitalia, dorsal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 329

SA

>|
Ry

1.0mm

FIGURE 18. Ceraeochrysa tenuicornis Adams and Penny, 1987. A. Head and prothorax, dorsal view; B. Wings; C. Apex of
male abdomen, lateral view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Gonapsis.

330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.0 mm

ISeeey T9+ect

FIGURE 19. Ceraeochrysa tucumana (Navas, 1919). A. Head and prothorax, dorsal view; B. Wings; C. Apex of male abdo-
men, lateral view; D. Sternite 8 + 9; E. Apex of male abdomen, dorsal view; F. Spermatheca; G. Subgenitale; H. Gonapsis; I.
Male genitalia, dorsal view; J. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 331

FIGURE 20. Ceraeochrysa dislepis, new species. A. Head and prothorax, dorsal view; B. Wings; C. Male genitalia, dorsal
view; D. Male genitalia, lateral view; E. Gonapsis; F. Apex of male abdomen, detail; G. Spermatheca.

332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.0 mm

0.5 mm

FIGURE 21. Ceraeochrysa dolichosvela, new species. A. Head and prothorax, dorsal view; B. Wings; C. Female genitalia.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 335

. s S80 A Y) i
Dies
scaly plate £~,7),lis

1.0 mm
1.0 mm

FIGURE 22. Ceraeochrysa squama, new species. A. Head and prothorax, dorsal view; B. Wings; C. Male genitalia, dorsal
view; D. Male genitalia, lateral view; E. Apex of male abdomen apex, lateral view; F. Male genitalia, caudal view; G. Gonapsis.

334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.0 mm 2.0 mm
E
D
ra acu
la

0.5 mm

FIGURE 23. Chrysoperla defreitasi Brooks, 1994. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D.
Tignum; E. Apex of male abdomen; F. Male genitalia, lateral view; G. Male genitalia, dorsal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 335

[TLE

1.0 mm

7 WN Pane!
H (sities)
FSby baer

SS

Se

0.5 mm

ger

0.5 mm

0.5 mm

FIGURE 24. Chrysoperla externa (Hagen, 1861). A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view; D.
Apex of male abdomen, lateral view; E. Tignum; F. Female genitalia; G. Subgentiale; H. Gonosaccus; I. Male genitalia, lateral
view; J. Male genitalia, dorsal view.

336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SSS eee

GSEGREEC
oe SS
TRESS

SS

<<
RRQ

0.5 cm

E tignum
S\

0.5 mm

gst
—_ arc
gc \ Sc
D tignum %

FIGURE 25. Chrysoperla raimundoi, new species. A. Head, frontal view; B. Wings; C. Apex of male abdomen; D. Male geni-

talia, dorsal view; E. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 337

T9+ect

0.5 mm

FIGURE 26. Chrysopodes (Chrysopodes) lineafrons Adams and Penny, 1987. A. Head and prothorax, dorsal view; B. Wings;
C. Head, frontal view; D. Apex of female abdomen, lateral view; E. Male sternite 8 + 9 apex; F. Male genitalia, dorsal view; G.
Male genitalia, lateral view; H. Spermatheca; I. Apex of male abdomen, lateral view.

338 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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—_—_s

0.5 mm

FIGURE 27. Chrysopodes (Chrysopodes) polygonica Adams and Penny, 1987. A. Head and prothorax, dorsal view; B.
Wings: C. Head, frontal view; D. Head, lateral view; E. Apex of forewing; F. Apex of female abdomen; lateral view; G. Apex of
male abdomen, lateral view; H. Male genitalia, dorsal view; I. Male genitalia, lateral view; J. Female genitalia; K. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 359

0.5 mm

FIGURE 28. Chrysopodes (Chrysopodes) spinella Adams and Penny, 1987. A. Head and prothorax, dorsal view; B. Wings: C.
Head, frontal view; D. Female genitalia.

340

C

FIGURE 29.
italia, latera

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

la g
D
\
gsc “ee =a
0.5 mm

Chrysopodes (Chrysopodes) adynatos, new species. A. Head and prothorax, dorsal view; B. Wings; C. Male gen-

view; D. Male genitalia, dorsal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 34]

SS LTTE
PEERS
LSS

FIGURE 30. Chrysopodes (Chrysopodes) copia, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Apex of male abdomen, lateral view; E. Subgenitale; F. Male genitalia, lateral view; G. Male genitalia, dorsal view; H.
Spermatheca.

342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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aX
TRS

F 0.5 mm arc

FIGURE 31. Chrysopodes (Chrysopodes) crocinus, new species. A. Wings; B. Apex of male abdomen, lateral view; C.
Encrassate median cell; D. Male genitalia, lateral view; E. Male genitalia, dorsal view; F. Spermatheca; G. Apex of female ab-
domen, lateral view; H. Subgenitale.

SS —

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 343

0.5 mm

FIGURE 32. Chrysopodes (Chrysopodes) delicata, new species. A. Head, frontal view; B. Wings; C. Head, lateral view; D.
Apex of male abdomen, lateral view; E. Mandibles; F. Male genitalia, dorsal view; G. Male genitalia, lateral view.

344 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.0 mm

FIGURE 33. Chrysopodes (Chrysopodes) elongata, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head,
frontal view; D. Female genitalia; E. Apex of female abdomen; F. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 345

FIGURE 34. Chrysopodes (Chrysopodes) nigropicta, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C.
male genitalia.

346 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SA

=<

1.0 mm C ag

FiGuRE 35. Chrysopodes (Neosuarius) divisa (Walker, 1853). A. Head and prothorax, dorsal view; B. Wings; C. Head, fron-
tal view; D, Apex of male abdomen; E. Female genitalia; F. Spermatheca; G. Male genitalia, dorsal view; H. Male genitalia, lat-

eral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 347

C
DONS »{ clypeusp
\ i
E sd
ay
vi

0.5 mm

FIGURE 36. Chrysopodes (Neosuarius) karinae Adams and Penny. A. Head and prothorax, dorsal view; B. Wings; C. Head,

frontal view; D. Apex of male abdomen, lateral view; E. Female genitalia; F. Apex of female abdomen, ventral view.

348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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TTI

Oe

5 6 A ST FE YR mse
a

0.5 mm

FIGURE 37. Plesiochrysa brasiliensis (Schneider, 1851). A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Apex of male abdomen, dorsal view; E. Apex of male abdomen, lateral view; F. Female genitalia; G. Male genitalia,
dorsal view; H. Male genitalia, lateral view; I. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 349

RAT Ik \

1.0 mm

FiGuRE 38. Plesiochrysa elongata (Navas, 1913). A. Head and prothorax, dorsal view; B. Wings; C. Head, lateral view.

350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.0 mm

FiGURE 39. Plesiochrysa alytos, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 351

T9+ect

“Blog 0.5mm

FIGURE 40. Leucochrysa (Leucochrysa) ampla (Walker, 1853). A. Wings; B. Head, frontal view; C. Head and prothorax, dor-
sal view; D. Abdomen, dorsal view; E. Female genitalia; F. Subgenitale.

352

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Volume 52, No. 19

=

blique
band
SQV
Ss
0.5cm
C T9+ect
16 T4 7 a 17

FiGuRE 41. Leucochrysa (Leucochrysa) boxi Navas, 1930. A. Head and prothorax, dorsal view; B. Wings; C. Abdomen, dor-
sal view; D. Apex of male abdomen, lateral view; E. Male genitalia dorsal; F. Sternite 8 + 9; G. Male genitalia, lateral view.

ee

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 353

7 [ [LLL

>
pes
S

SR eS
me? URS
Sa

Tan @melal TS
SEEAAINAWWNER

0.5mm 2.5mm

FIGURE 42. Leucochrysa (Leucochrysa) pretiosa (Banks, 1910). A. Head, frontal view; B. Wings; C. Apex of abdomen, lat-
eral view; D. Male genitalia, dorsal view; E. Head and prothorax, dorsal view; F. Female genitalia; G. Male genitalia, lateral
view; H. Subgenitale, dorsal view; I. Subgenitale, lateral view; J. Abdomen, dorsal view.

354 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Sa LS
URS

|] [LL LTEEE zzz
quae
cues

2.5mm

T8
. ge

FiGURE 43. Leucochrysa (Leucochrysa) varia (Schneider, 1851). A. Wings; B. Head and pro- and mesothorax, dorsal view;
C. Abdomen, dorsal view; D. Head, frontal view; E. Female genitalia; F. Subgenitale; G. Male genitalia, dorsal view; H. Male
genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 355

<—>>

&

Bac

|B HAS
[is

I EAR SS
IEICE
ZZ ees
1 ZW OK
Lh

ih

0.5mm

FiGuRE 44. Leucochrysa (Leucochrysa) walkerina Navas, 1913. A. Wings; B. Head, pro- and mesothorax, dorsal view; C.
Head, frontal view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view.

356 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES |
Volume 52, No. 19

| [LLL eae

oats SHEL ~

2.5mm

FIGURE 45, Leucochrysa (Leucochrysa) bruneolus, new species. A. Wings; B.
Male genitalia, dorsal view; D. Male genitalia, lateral view; E. Male genitalia, caudal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE)

1.0 mm

RE rysa ( chrysa) W

358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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TT PPR Z2

Bee
NI

FIGURE 47. Leucochrysa (Nodita) camposi Navas, 1933. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Male genitalia, ventral view; G. Meso- and metanotum;
H. Apex of abdomen, lateral view; I. Abdomen, dorsal view; J. Subgenitale; K. Female genitalia.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 359

gen

1.0mm

FiGuRE48. Leucochrysa (Nodita) clepsydra Banks, 1918. A. Head and prothorax, dorsal view; B. Forewing; C. Head, frontal
view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Male genitalia, caudal view.

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FiGuRE 49. Leucochrysa (Nodita) cruentata (Schneider, 1851). A. Head, pro- and mesothorax, dorsal view; B. Wings; C.
Head, frontal view; D. Abdomen, dorsal view; E. Apex of male abdomen, lateral view; F. Male genitalia, dorsal view; G. Male
genitalia, lateral view; H. Male genitalia, caudal view; I. Spermatheca; J. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 361

0.5 cm

0.5 mm

I

+
aN
ss) ventral lobe

FIGURE 50. Leucochrysa (Nodita) gossei (Kimmins, 1940). A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Head,
frontal view; D. Abdomen, dorsal view; E. Male genitalia, dorso-caudal view; F. Male genitalia, dorsal view; G. Male genitalia,
lateral view; H. Female genitalia; I. Subgenitale.

362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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< T9+ect
~\ a

1.0mm

FIGURE 51. Leucochrysa (Nodita) heriocles Banks, 1944. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Head,
frontal view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Male genitalia, caudal view; G. Spermatheca; H.
Abdomen, dorsal view; I. Apex of male abdomen, lateral view; J. Sternite 8 + 9.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 363

fir,
=e

y
== basal
iS a stalk

BESS

1.0mm

FIGURE 52. Leucochrysa (Nodita) intermedia (Schneider, 1851). A. Head and prothorax, dorsal view; B. Wings; C. Abdo-
men, dorsal view; D. Female genitalia; E. Subgenitale.

364 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 53. Leucochrysa (Nodita) lancala Banks, 1944. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Male genitalia, caudal view; G. Abdomen, dorsal view;
H. Apex of male abdomen, lateral view; I. Subgenitale; J. Spermatheca.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 365

a= =. .

FiGurRE 54. Leucochrysa (Nodita) lateralis Navas, 1913. A. Head, thorax and abdomen, dorsal view; B. Wings; C. Head,
frontal view; D. Spermatheca; E. Subgenitale.

366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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» St
RAS

imc

SS Snes ce
RESTA RSA

0.5 cm

FIGURE 55. Leucochrysa (Nodita) marginalis (Banks, 1915). A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Apex of male abdomen, lateral view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Male genitalia,
caudal view; H. Subgenitale; I. Female genitalia.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 367

0.5 mm

FIGURE 56. Leucochrysa (Nodita) marquezi Navas, 1917. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Head, lateral view; E. Abdomen, dorsal view; F. Spermatheca; G. Female genitalia; H. Male genitalia, lateral view; I.
Male genitalia, dorsal view; J. Subgenitale.

368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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eas
—e

FIGURE 57. Leucochrysa (Nodita) melanocera Navas, 1916. A. Head, prothorax and abdomen, dorsal view; B. Wings; C. Fe-
male genitalia; D. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 369

<< e a.

LJ N
REQ
al

iS
Waren
StS

1.0 mm

1.0mm

FIGURE 58. Leucochrysa (Nodita) rodriguezi (Navas, 1913) A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Abdo-
men, dorsal view; D. Head, frontal view; E. Apex of male abdomen, lateral view; F. Sternite 8 + 9; G. Male genitalia, dorsal
view; H. Male genitalia, lateral view.

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1.0mm

FIGURE 59. Leucochrysa (Nodita) affinis, new species. A. Head and prothorax, dorsal view; B. Wings; C. Abdomen, dorsal
view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) S71

1.0mm

FIGURE 60. Leucochrysa (Nodita) barrei, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Head, fron-
tal view; D. Head, lateral view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

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Fe
Sse a

T9+ect

‘My, bie

FIGURE 61. Leucochrysa (Nodita) confusa, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Abdomen,
dorsal view; D. Male genitalia, dorsal view; E. Male genitalia, lateral view; F. Male genitalia, caudal view; G. Abdomen, dorsal
view; H. Apex of male abdomen, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 373

B ] (GE Bless = ol

ES

Ww

ey:

FIGURE 62. Leucochrysa (Nodita) cornuta, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal
view; D. Male genitalia, lateral view; E. Male genitalia, dorsal view; F. Male genitalia, caudal view.

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Mibeasass

APPL EL
See.

0.5mm

FIGURE 63. Leucochrysa (Nodita) forciformis, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Abdo-
men, dorsal view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Female genitalia.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 375

Ree NN
RRS

FIGURE 64. Leucochrysa (Nodita) furcata, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Abdomen,
dorsal view; D. Apex of male abdomen; E. Sternite 8 + 9; F. Head, frontal view; G. Male genitalia, dorsal view; H. Male genita-
lia, lateral view; I. Female genitalia; J. Spermatheca.

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FiGURE 65. Leucochrysa (Nodita) guataparensis, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings, C. Ab-
domen, dorsal view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Female genitalia;
I. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) B77

FIGURE 66. Leucochrysa (Nodita) ictericus, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Head,
frontal view; D. Head, lateral view; E. Male genitalia, dorsal view, F. Male genitalia, lateral view; G. Male genitalia, caudal
view; H. Apex of female abdomen, lateral view; I; Spermatheca; J. Subgenitale.

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lL LJ LLLA ae

a SNe
(es \\ SS arc
S Sok Stk

SS,

S8+9

FIGURE 67. Leucochrysa (Nodita) incognita, new species A. Wings; B. Apex of male abdomen, lateral view; C. Head, frontal
view; D. Male genitalia, dorsal, lateral and caudal view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE)

Zl el Oe eee
=e

Bee eee
REI

PERRRRESA

TERS

Sp

Sa
pea

= (i

0.5mm

1.0mm

FIGURE 68. Leucochrysa (Nodita) interata, new species. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Head,
ontal view; D. Head, lateral view; E. Abdomen, dorsal view; F. Spermatheca; G. Subgenitale.

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FIGURE 69. Leucochrysa (Nodita) lineata, new species. A. Pro-, meso- and metanota; B. Wings; C. Head, frontal view; D.
Abdomen, dorsal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Gonosetae.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 381

1.0mm

0.5 mm

FIGURE 70. Leucochrysa (Nodita) maculata new species. A. Head and thorax, dorsal view; B. Wings; C. Spermatheca; D.
Head, frontal view.

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FIGURE 71. Leucochrysa (Nodita) michelini, new species. A.Thorax, dorsal view; B. Wings; C. Abdomen, dorsal view; D.
Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 383

T9+ect

1.0mm

1.0mm

FIGURE 72. Leucochrysa (Nodita) parallela new species A. Head and thorax, dorsal view; B. Wings; C. Apex of male abdo-
men, lateral view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

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FIGURE is ips ochrysa (No we g)re retusa. A. Head, pro- and mesothorax, dorsal view; B. Wings; C. Male genitalia, dorsal
vi oe lia, lateral v

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE)

385

FIGURE 74. Leucochrysa (Nodita) robusta, new species. A. Head and prothorax, dorsal view; B. Wings; C. Abdomen, dorsal
view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Female genitalia; H. Subgenitale.

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0.5mm

1.0mm

FIGURE 75. Leucochrysa (Nodita) santni, new species. A. Head and prothorax, dorsal view; B. Wings; C. Abdomen, dorsal
view; D. Head, frontal view; E. Male genitalia, dorsal view; F. Male genitalia, lateral view; G. Subgenitale.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 387

0.5mm

FIGURE 76. Leucochrysa (Nodita) scomparini, new species. A. Head and thorax, dorsal view; B. Wings; C. Head, frontal
view; D. Subgenitale; E. Female genitalia.

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FIGURE 77. Leucochrysa (Nodita) squamisetosa, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, fron-
tal view; D. Abdomen, dorsal view; E. Male genitalia, dorso-caudal view; F. Male genitalia, dorsal view; G. Male genitalia, lat-
eral view.

i
|

|
J

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 389

FIGURE 78. Leucochrysa (Nodita) tabacinus, new species A. Head and thorax, dorsal vie
D. Apex of female abdomen, lateral view; E. Male genitalia, dorsal view; F. Male genitali
view; H. Female genitalia; I. Subgenitale.

w; B. Wings; C. Head, frontal view;
a, lateral view; G. Abdomen, dorsal

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=o Sennleisonc
Res

=
Lx Ip

gen

0.5mm

FIGURE 79. Leucochrysa (Nodita) tenuis, new species. A. Head and prothorax, dorsal view; B. Wings; C. Head, frontal view;
D. Male genitalia, dorsal view; E. Male genitalia, lateral view.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 39]

FiGure 80. Leucochrysa (Nodita) vignisi, new species. A. Head and thorax, dorsal view; B. Wings; C. Head, frontal view; D.,
E. Male genitalia, dorsal view; F. Male genitalia, lateral view.

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0.5mm

FIGURE 81. Leucochrysa (Nodita) vittata, new species. A. Head and thorax, dorsal view; B. Wings; C. Abdomen, dorsal
view; D. Head, frontal view; E. Spermatheca; F. Subgenitale, dorsal view; G. Subgenitale, lateral view.

ri - 20 we —————

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 393

ACKNOWLEDGMENTS

This project could not have been accomplished without the extensive help provided by a series of
notebooks of detailed drawings and notes of type specimens developed by Phillip A. Adams. These
notes cover almost all existing types of Neotropical chrysopids in European and North American col-
lections. However, any errors in interpretation of these notes are our responsibility. We gratefully ac-
knowledge a grant from the Ernst Mayr Fund at the Museum of Comparative Zoology, Harvard
University for allowing the junior author to study types and determined Neotropical material in that
museum. This work was also supported by Fundacgao de Amparo a Pesquisa do Estado de Sao Paulo —
FAPESP, Project No. 99/05245-2.

LITERATURE CITED

ADAMS, P. A. 1977. Taxonomy of United States Leucochrysa (Neuroptera: Chrysopidae). Psyche, Camb.
84(1):92—102.

. 1978. Zoogeography of New World Chrysopidae, a progress report. Folia Entomologica Mexicana

39/40:210-211.

. 1982a. Ceraeochrysa, a new genus of Chrysopinae (Neuroptera) (Studies in New World Chrysopidae,

Part II). Neuroptera International 2:69—75, 12 figs.

. 1982b. Plesiochrysa, a new subgenus of Chrysopa (Neuroptera) (Studies in New World Chrysopidae,
Part I). Neuroptera International 2:27—32, 13 figs.

ADAMS, P. A. ANDN. D. PENNY. 1987. Neuroptera of the Amazon Basin. Part | 1a. Introduction and Chrysopini.
Acta Amazonica 15:413-479. 213 + 29 figs., 1 table.

BANKS, N. 1910. New South American neuropteroid insects. Proceedings of the Entomological Society of Wash-
ington 12:146—160.

. 1915. New neuropteroid insects, native and exotic. Proceedings of the Academy of Natural Sciences of

Philadelphia 66:608-632, | pl. with 26 figs.

. 1918. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology 62:1—22, 2 pls. with

26 figs.

. 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology 64:297-362, 7 pls.

with 110 figs.

. 1944. Neuroptera of northern South America. Part III. Chrysopidae. Boletin de Entomologia

Venezolana 3:1—34.

. 1945. A review of the Chrysopidae (Nothochrysidae) of Central America. Psyche, Camb. 52:139-174.

. 1948. Chrysopidae (Nothochrysidae) collected in Mexico by Dr. A. Dampf (Neuroptera). Psyche,
Camb. 55:151—177, 3 pls. with 37 figs.

BROOKS, S. J. 1994. A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera:
Chrysopidae). Bulletin of the British Museum of Natural History, Entomology Series 63(2):137-210.

BROOKS, S. J. AND P. C. BARNARD. 1990. The green lacewings of the world: a generic review (Neuroptera:
Chrysopidae). Bulletin of the British Museum of Natural History, Entomology Series 59:1 17-286, 578
figs., 1 table.

GERSTAECKER, A. 1888. Weitere Beitrage zur artenkenntniss der Neuroptera Megaloptera. Mitteilungen des
Naturwissenschaftlichen Vereins fiir Neu- Vorpommern und Rugen in Greifswald 19:89—130.

HAGEN, H. 1861. Synopsis of the Neuroptera of North America, with a list of the South American species. Smith-
sonian Miscellaneous Collections 49(1):xx + 1-347.

KiImMINns, D. E. 1940. Notes on some types of Chrysopidae (Neuroptera) in the British Museum Collections. An-
nals and Magazine of Natural History (11)5:442-449.

LOPEZ-ARROYO, J. I., C. A. TAUBER AND M. J. TAUBER. 1999. Comparative life-histories of the predators
Ceraeochrysa cincta, C. cubana, and C. smithi (Neuroptera: Chrysopidae). Annals of the Entomological
Society of America 92:208—217.

MATSUDA, M. 1928. Observations on Chrysopa vulgaris Schn. var. ampingensis, Petersen. Transactions of the
Natural History Society of Formosa 18:97—114. [in Japanese]

394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 19

MCLACHLAN, R. 1868. New genera and species, &c., of neuropterous insects; and a revision of Mr. F. Walker’s
British Museum Catalogue of Neuroptera, part ii. (1853), as far as the end of the genus Myrmeleon. Journal
of the Linnean Society of London, Zoology 9:230—281, | pl. with 3 figs.

MINISTERIO DE AGRICULTURA. 1999. www.minagric.br.

NavAs, L. 1911. Chrysopides nouveaux (Ins. Neur.). Annales de la Société Scientifique de Bruxelles 35 (pt.
2):266—282, 7 figs.

. 1913a. Crisopidos sudamericanos. Brotéria (Zooldgica) 11:73-104, 149-168. 8 and 5 figs.

. 1913b. Névroptéres nouveaux de l’Amérique du Nord [I]. Entomologische Zeitschrift, Frankfurt am

Main 27:19-20, 2 figs.

. 1913c. Les Chrysopides (Ins. Névr.) du Musée de Londres [la]. Annales de la Société Scientifique de

Bruxelles 37 (pt. 2):292—330, 19 figs.

. 1914. Les Chrysopides (Ins. Névr.) du Musée de Londres [1b]. Annales de la Société Scientifique de

Bruxelles 38 (pt. 2)73—114, 15 figs.

. 1915. Neurdpteros nuevos 0 poco conocidos. (Sexta [VI] serie). Memorias de la real Academia de

Ciencias y Artes de Barcelona (3)12:119—136, 9 figs.

. 1916. Neurdpteros sudamericanos. (Trecera [III] serie). Neurdpteros del Brasil recogidos por el R. P.

Joaquin da Silva Tavares S. J. Brotéria (Zoolégica) 14:14—35, 15 figs.

. 1919. Algunos insectos Neurdpteros de la Republica Argentina. (Trecera [III] serie). Revista de la Real

Academia de Ciencias Exactas Fisicas y Naturales de Madrid 17:287-—305, 6 figs.

. 1920. Insectos Sudamericanos (3a serie). Anales de la Sociedad Cientifica Argentina 90:52-72, 11 figs.

. 1924a. Insectos de la Argentina y Chile. Estudios. Revista Mensual (Academia literaria del Plata, Bue-

nos Aires) 22:358—368. 4 figs.

. 1924b. Crisopidos (Neur.) de Cuba. Boletin de la Sociedad Entomologica de Espana 7:51—S3.

. 1926. Algunos insectos del Brasil (3.a serie) [IIIb]. Brotéria (Zoologica) 23:5—15, 5 figs.

. 1930. Insectos de la Argentina. (Sexta [VI] serie). Revista de la Sociedad Entomologica Argentina

3:125-—132, 5 figs.

. 1933. Insectos suramericanos. (Sexta [VI] serie). Revista de la Real Academia de Ciencias Exactas
Fisicas y Naturales de Madrid 29:191—198, 7 figs.

NEUMARK, A. 1952. Chrysopa carnea Steph. and its enemies in Israel. Forest Research Station, Ilanoth |:vii +
1-127, 68 figs.

PENNY, N. D. 1978. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, ilhas
Caraibas e América do Sul. Acta Amazonica 7(4)(Suplemento): 1-61.

. 1997. Four new species of Costa Rican Ceraeochrysa (Neuroptera: Chrysopidae). Pan-Pacific Ento-

mologist 73(2):61—69.

. 1998. New Chrysopinae from Costa Rica (Neuroptera: Chrysopidae). Journal of Neuropterology

1:55—78. 68 figs.

. 2001. New species of Chrysopinae (Neuroptera: Chrysopidae) from Costa Rica, with selected taxo-
nomic notes and a neotype designation. Entomological News 112(1):1—14.

PENNY, N. D., P. A. ADAMS, AND L. A. STANGE. 1997. Species catalog of the Neuroptera, Megaloptera, and
Raphidioptera of America north of Mexico. Proceedings of the California Academy of Sciences
50(3):39-114.

SCHNEIDER, W. G. 1851. Symbolae ad monographiam generis Chrysopae, Leach. Hirt, Vratislaviae. 178 pp., 60
pls.

SMITH, R. C. 1921. A study of the biology of the Chrysopidae. Annals of the Entomological Society of America
14:27-35.

. 1922. The biology of the Chrysopidae. Memoirs of the Cornell University Agricultural Experiment sta-
tion 58:1287—1372, 10 text figs. and 14 pls. with 122 figs.

STEINMANN, H. 1964. The Chrysopa species (Neuroptera) of Hungary. Annales Historico-Naturales Musei
Nationalis Hungarici (Zoologica) 56:257—266, | fig.

TAUBER, C. A. AND T. DE LEON. 2001. Systematics of green lacewings (Neuroptera: Chrysopidae): Larvae of
Ceraeochrysa from Mexico. 94(2):197—209.

TAUBER, C, A., T. DE LEON, N. D. PENNY, AND M. J. TAUBER. 2000. The Genus Ceraeochrysa (Neuroptera:
Chrysopidae) of America north of Mexico: Larvae, adults, and comparative biology. Annals of the Entomo-
logical Society of America 93(6):1195—1221.

DE FREITAS AND PENNY: GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE) 395

TAUBER, M. J.,C. A. TAUBER, K. M. DANNE, AND K. S. HAGEN. 2000. Commercialization of predators: Recent
lessons from green lacewings (Neuroptera: Chrysopidae: Chrysoperla). American Entomologist
46(1):26-38.

TJEDER, B. 1966. Neuroptera-Planipennis. The Lace-wings of Southern Africa. 5. Family Chrysopidae. Pp.
228-534 in South American Animal Life. B. Hanstrém, P. Brinck, and G. Rudebeck, eds. Vol. 12. Swedish
Natural Science Research Council, Stockholm.

WALKER, F. 1853. List [Catalogue] of the specimens of neuropterous insects in the collection of the British Mu-
seums. Part II. (Sialidaes - Nemopterides). British Museum [Natural History], London. Pp. 193-476.
WITHYCOMBE, C. L. 1923. Notes on the biology of some British Neuroptera (Planipennis). Transactions of the

Entomological Society of London 1922:501—594, 6 pls. with 72 figs.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA. Sate OF. Se

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October 26, 2001
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Volume 52, No. 20, pp. 397-405, 5 figs.

A New Lycodon (Serpentes: Colubridae) from Northeast India
and Myanmar (Burma)

by

Joseph B. Slowinski!, Samraat S. Pawar?, Htun Win, Thin Thin?, Sai Wanna Gyi’,
San Lwin Oo, and Hla Tun?

1 Department of Herpetology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118
2Centre for Ecological Research and Conservation, 3076/5 IV Cross, Gokulam Park,
Mysore 570 002, India; Tel: 91- 821- 515-601; email samraat@ncf-india.org
3 Nature and Wildlife Conservation Division, Forest Department, Ministry of Forestry,
Bayintnaung Road, West Gyogone, Insein, Yangon, Myanmar

A new species of the colubrid snake genus Lycodon is described from western Myanmar
and Northeast India. Morphologically, the new species is most similar to L. /aoensis, but is
distinguishable from it by its brownish black and white dorsal coloration (vs. brownish
black and yellow in L. /aoensis) and the absence of a well-developed nape band (present in
L. laoensis). The new species increases the diversity of Lycodon to four in Myanmar and to
five in Northeast India.

Lycodon H. Boie in Fitzinger (1826) is a genus of small colubrid snakes characterized by an
arched maxillary generally with three sets of teeth separated by two diastemata, a dorsoventrally com-
pressed head, and a vertically elliptical pupil (Smith 1943; Taylor 1965; Leviton 1965). Approxi-
mately 25 species occur in Asia (Lanza 1999). Recently, herpetofaunal surveys in Myanmar and
Northeast India (by Slowinski, Htun Win, Thin Thin, Sai Wanna Gyi, San Lwin Oo, and Hla Tun in
Myanmar; Pawar in Northeast India) yielded specimens of Lycodon from western Myanmar and
Northeast India that represent a new species. Four other species, viz., L. aulicus/capucinus, L.
fasciatus, L. kundui, L. laoensis, and L. jara, occur in Myanmar and Northeast India (Smith 1943;
Dowling and Jenner 1988).

All specimens were hand collected, euthanized, fixed in 10% buffered formalin and later trans-
ferred to 70% ethanol. Comparative material was examined at the CAS (California Academy of Sci-
ences), BMNH (Bombay Museum of Natural History), and ZSI (Zoological Survey of India). In
addition, information was collected from private collections in India and field stations in various pro-
tected areas in Northeast India.

1 Dr. Joseph Slowinski died in Myanmar on September 12, 2001, while leading a biological expedition to the
extreme northeast of that country. He will be deeply missed by his academic colleagues, his students and friends,
and by his many co-workers in Myanmar.

397

398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 20

SPECIES DESCRIPTION
Lycodon zawi sp. nov.
Zaw’s wolf snake

MATERIAL EXAMINED. — HOLOTYPE: CAS 210323 (Fig. 1), a male from Alaungdaw Katapha
National Park (AKNP) (22°19’'N, 94°29’E), Sagaing Division, Myanmar, collected 17 July, 1999, by
Htun Win, Thin Thin, K. Wiseman, J. Lovette, and J. Vindum. PARATYPES: CAS 210223, a male from
AKNP (22°19’N, 94°24’E), Sagaing Division, Myanmar, collected 10 July, 1999, by J. Slowinski, K.
Wiseman, J. Lovette, and J. Vindum; CAS 215494, a male from AKNP (22°19'N, 94°24’E), Sagaing
Division, Myanmar, collected 30 May, 2000, by Htun Win, Thin Thin, San Lwin Oo, Sai Wanna Gyi,
and Hla Tun; CAS 215570, a male from AKNP (22°19’N, 94°29’E), Sagaing Division, Myanmar, col-
lected 12 June, 2000, by Htun Win, Thin Thin, and San Lwin Oo; CAS 215599, a male from AKNP
(22°19'N, 94°29’E), Sagaing Division, Myanmar, collected 14 June, 2000, by Htun Win, Thin Thin,
and San Lwin Oo; CAS 216505, a male from the Gwa Township (17°39'N, 94°39’E), Rakhine State,
Myanmar, collected 29 November, 2000, by J. Slowinski, Htun Win, and Hla Tun. ZSI 25346, a male
from Ngengpui Wildlife Sanctuary (NgWS; 22°29’'N, 92°48’E), Mizoram, Northeast India, collected
17 April, 1999, by S. Pawar; ZSI 25347, a male from Nongkhyllem Wildlife Sanctuary (NWS;
25°56'N, 91°31'E), Meghalaya, Northeast India, collected 8 May, 2000, by M. F. Ahmed; ZSI 25348,
a female from Garbhanga Reserve Forest (GRF; 26°09’N, 91°33’E), Assam, Northeast India, col-
lected 30 March, 1998, by S. Sengupta. ADDITIONAL SPECIMENS: Two more specimens, a male near
Kaifung (23°39'N, 92°57’E), North Mizoram, Northeast India, and another male from Balphakram
Tiger Reserve (BTR; 25°30'N, 90°45’E), Meghalaya, Northeast India, not housed in a permanent de-
pository.

DIAGNOSIS. — Lycodon zawi differs from other Lycodon of the Asian mainland by the following
combination of character states: 17 dorsal scale rows at mid-body, preocular scale present (Fig. 1),
loreal scale not in contact with internasal (Fig. |), anal scale divided, poorly-developed white
crossbands on a brownish black dorsum, and without a well-developed nape band (Fig. 1). In terms of
scale characteristics, L. zawi is similar to L. laoensis (Figs. 2-4), known from Northeast India, China,
Malaysia, Thailand, Laos, Vietnam, and Cambodia (Lanza 1999), but differs by its brownish black
and white dorsal coloration (brownish black and yellow in L. laoensis), the poorly-developed light
crossbands (well-developed in L. /aoensis), and the lack of a nape band. Lycodon zawi differs from
other Myanmar and Northeast India congeners as follows: from L. aulicus/capucinus (Figs. 2—4) in
lacking a nape band, in lacking a sharply defined white lip margin, and in having a loreal scale that
does not contact the internasal scale; from L. kundui in lacking a nape band, having 17 mid-body scale
rows (15 in L. kundui), and in having 8 or 9 supralabials (7 in L. kundui); from L. fasciatus (Figs. 2-4)
in lacking loreal contact with the eye, in lacking well-developed light crossbands, and in having
smooth dorsal scales; from L. jara (Figs. 2—3) in having light bands and in having a loreal scale that
does not contact the internasal scale. Lycodon zawi resembles L. travancoricus (Western Ghats and
southern Pakistan) and L. tiwarii (Andaman and Nicobar Islands; Biswas and Sanyal 1965) in
scalation. From L. travancoricus, L. zawi differs by its divided anal and weakly developed light bands
(L. travancoricus has well-developed yellow bands which bifurcate on the sides). Lycodon zawi dif-
fers from L. tiwarii in having white bands on a dark dorsum (white reticulations on a dark dorsum in L.
tiwarii).

DESCRIPTION OF HOLOTYPE (Adult male). — Body dimensions: SVL 395 mm; tail length
85 mm; total length 480 mm. Body scalation: 183 ventrals; 45 subcaudals; 17-17-15 dorsal scale
rows. Head scalation: loreal well separated from internasal and from the eye border by the preocular
and 3rd supralabial; 8 supralabials, 3rd, 4th, and Sth touching eye; | postocular; 2+3 temporals; 9/10
infralabials, Ist to Sth infralabials contacting chin shields.

SLOWINSKI ET AL.: LYCODON (SERPENTES: COLUBRIDAE) Beis)

FiGure |. The holotype (CAS 210323) of Lycodon zawi.

400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 20

FIGURE 2. Dorsal views of heads of (top to bottom) Lycodon zawi (CAS 210323), Lycodon fasciatus (CAS 55147) from west-
ern China, L. aulicus (CAS 216278) from Mandalay Division, Myanmar, L. /aoensis (CAS 73679) from Thailand, and L. jara
(CAS 12395) from Assam, India.

SLOWINSKI ET AL.: LYCODON (SERPENTES: COLUBRIDAE) 401

FicurE 3. Lateral views of heads of (top to bottom; same snakes as in Fig. 2) Lycodon zawi, L. fasciatus, L. aulicus, L.
laoensis, and L. jara.

402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 20

The maxilla (right maxilla examined) is arched and similar in form to that illustrated by Smith
(1943, fig. 88) for L. aulicus. Two diastemata separate the maxillary teeth into three groups. The ante-
rior group is composed of three small teeth followed posteriorly by two enlarged teeth; the middle
group is composed of five equal teeth; the posterior group is composed of two enlarged teeth.

Everted hemipenis extending to 8th subcaudal; unforked; distal half characterized by longitudi-
nal flounces perpendicular to the long axis of organ; proximal half of organ with large spines; sulcus
single, terminating in expanded trough at tip.

Coloration in life (same as in preservative): brownish black dorsally with irregularly shaped
white transverse bands, the bands well developed anteriorly (one scale row in width) but fading poste-
riorly; faint, diffuse light band on nape; head dark brown dorsally, fading to light brown on lips; tail
uniform brown dorsally without white bands; venter cream with dark lateral corners on each ventral
scale.

VARIATION. — Four of six Myanmar specimens have 8 supralabials on both sides, but two have
9/8 supralabials. All five Northeast India specimens have 9 supralabials on both sides. Five of the six
Myanmar specimens have 9 infralabials on both sides, but the holotype has 9/10. All five Northeast
India specimens have 10 infralabials. The holotype, a paratype from Northeast India (ZSI 25348), and
two other specimens from Northeast India have 2+3 temporals on both sides. Another paratype from
Northeast India (ZSI 25347) has 2+3 temporals on the left and 3+4 on the right. The third paratype
from Northeast India (ZSI 25346) has 2+1 temporals on the left and 2+2 on the right. All other speci-
mens, from Myanmar, have 1+2+3 temporals. The holotype and two other Myanmar specimens (CAS
210223, 1/1 postocular; CAS 216505, 2/1 postocular) have one postocular; all other specimens have
two postoculars. Ventrals in males range from 179 to 186 in the Myanmar specimens and 190 to 194 in
the Northeast India specimens; the single female specimen (ZSI 25348) from India has 207 ventrals;
subcaudals in males range from 45 to 67 in the Myanmar specimens and 70 to 75 in the Northeast India
specimens; the single female from India has an incomplete tail.

The general color pattern is identical in all specimens, except for some white bands on the tail in
several specimens. Three Northeast India specimens have a faint spot on the nape. The ventral color-
ation of the female paratype (ZSI 25348) consists of erratically distributed dark patches on the ventral
scales, not seen in the others.

ETYMOLOGY. — The specific name is a patronym in the genitive singular, honoring U Khin
Maung Zaw, Director of the Myanmar Nature and Wildlife Conservation Division, who has provided
critical assistance to our survey of the herpetofauna of Myanmar.

DISTRIBUTION AND NATURAL HISTORY. — In Myanmar, Lycodon zawi is currently known from
two localities in the west (Fig. 5): five specimens collected from AKNP in the Sagaing Division, and
one specimen collected from a locality on the western slope of the southern Rakhine (Arakan) Moun-
tains, over 470 km south of the first locality. Alaungdaw Katapha National Park is in the Sagaing Di-
vision in west-central Myanmar, approximately 160 km west of Mandalay. This is an area of low
mountains with a maximum elevation of 1000 m. Rainfall at AKNP averages 1500 mm per year (Tun
Nyo 1997), and the park consists of a mosaic of deciduous forest types, from closed canopy moist de-
ciduous forest to indaing, a savanna of stunted dipterocarp trees. The single Rakhine Yoma specimen
came from the forests near Gwa in the Rakhine State, approximately 170 km west of Yangon (Ran-
goon). Rainfall in the southern Rakhine mountains is quite high, averaging over 5000 mm per year
(data from Myanmar government). Owing to this, the habitat is quite different from that in AKNP. Or-
iginally evergreen forest (Stamp 1924, 1930), logging and shifting cultivation has reduced most of the
forest to extensive bamboo stands (Collins et al. 1991).

In Northeast India, L. zawi has been collected at five localities (Fig. 5). All localities are in low to
mid-elevation hill tracts receiving medium to high precipitation (above 2000 mm per year); all sites
harbor, or formerly harbored, low to mid-elevation moist tropical evergreen to semi-evergreen forest.
Four of the five localities, viz., NgWS, NWS, BTR, and GRF, are protected. As in Myanmar, habitat

SLOWINSKI ET AL.: LYCODON (SERPENTES: COLUBRIDAE) 403

FiGure 4. Anterior bodies of (top to bottom; same snakes as in Fig. 2) Lycodon fasciatus, L. aulicus, and L. laoensis.

alteration due to slash-and-burn cultivation is a major problem for habitat conservation in Northeast
India (Ramakrishnan 1992). In fact, the specimen (ZSI 25346) from NgWS was caught while fleeing a
burning slash-and-burn plot near the sanctuary boundary.

All evidence suggests that L. zawi is a nocturnal species, apparently preferring riparian forests.
Most specimens were found active at night along streams at elevations less than 500 m. Although
there are extensive tracts of bamboo in Northeast India-Myanmar, no specimens were found in bam-
boo habitats.

In both Myanmar localities, as well as most of the localities in Northeast India, L. zawi is
sympatric with L. aulicus and/or L. fasciatus. All localities of L. zawi from Northeast India are south
of the Brahmaputra river, which is an important biogeographical barrier in that region (Mani 1974;
Ripley and Beehler 1990). Despite recent surveys (Pawar, unpublished data), no specimens have been
obtained north of the Brahmaputra river, including the eastern Himalayas. If found there, L. zawi may
be sympatric with as many as three other species of Lycodon: L. jara, L. aulicus, and L. laoensis.

Three specimens of Lycodon zawi from Myanmar had prey items in their alimentary tracts: CAS
210223 contained the partially digested head of a small skink; CAS 215494 contained the rear torso
and tail of a small skink; CAS 215599 contained a hind limb and the tail of a small skink. In each case,
the skink appears to be Sphenomorphus maculatus, which is common along streams in Myanmar. One
specimen (ZSI 25346) kept by Pawar in captivity for 28 days fed on geckos (Hemidactylus frenatus
and H. garnoti).

DISCUSSION
Lycodon zawi seems to be common where it occurs. Its recent discovery is not surprising, because

western Myanmar and Northeast India remain very poorly surveyed. Recent surveys of this region by
the authors have brought to light new records and species for the region (Slowinski and Wuster 2000;

404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 20

LEU
_|CAS 215598) Zod a

“Icas 210223

Lycodon zawi
1999-2000

® Collection locality

California Academy of Sciences
Department of Herpetology

100 200

87° 92° 97° 102°
FiGureE 5. Distribution of Lycodon zawi in Myanmar and India.

Pawar and Biswas 2001; Pawar and Choudhury 2000). It is also possible that specimens of this species
exist in local museums in India, perhaps misidentified as L. aulicus or L. laoensis, to which the new
species bears superficial resemblance. In general, L. zawi is easily differentiated by its combination of
morphology and color pattern. It is apparent from the above description that there is some variation
both within and between Northeast India and Myanmar samples. The apparent differences between
the ventral scale counts of the Myanmar and Northeast India populations suggests geographic
differentiation.

ADDITIONAL MATERIAL EXAMINED

Lycodon laoensis.—CAS 73679 (Thailand), CAS-SU 8523 (Penang, Malaysia), CAS 15966
(Kerala, India).

Lycodon aulicus.—CAS 216278 (Mandalay Division, Myanmar), CAS 215387 (Sagaing Divi-
sion, Myanmar), CAS 215396 (Sagaing Division, Myanmar), CAS 215422 (Sagaing Division,
Myanmar).

Lycodon fasciatus —CAS 55147 (China), CAS 172715 (Chiang Mai Province, Thailand).

Lycodon jara.—CAS 17210 (Orissa, India), CAS-SU 12395 (Assam, India).

Lycodon tiwarii.—ZSI 20849 (no locality available), CAS 20851 (no locality available).

Lycodon travancoricus.—CAS-SU 15967 (Kerala, India).

SLOWINSKI ET AL.: LYCODON (SERPENTES: COLUBRIDAE) 405

ACKNOWLEDGMENTS

Financial support for Slowinski, Htun Win, Thin Thin, Sai Wanna Gyi, San Lwin Oo, and Hla
Tun’s field work in Myanmar was provided by funds from NSF grant DEB-9971861 and from the
California Academy of Sciences. U Uga and U Khin Maung Zaw, past and present directors of the
Myanmar Nature and Wildlife Conservation Division, respectively, have encouraged and assisted our
work in Myanmar. For assistance in the field, we thank J. Lovett, D. Lin, H. Robeck, J. Vindum, and
K. Wiseman. For other forms of assistance, we thank D. Lin. Surveys in Northeast India were sup-
ported by the Wildlife Institute of India, Bombay Natural History Society, and the Chicago Zoological
Society. We thank S. Sengupta and M. F. Ahmed for providing specimens and data from Northeast In-
dia. We also thank the Forest Department of Mizoram for providing Pawar with the necessary permits,
and their invaluable assistance in the field. Alan Leviton, D. Lin, and M. Koo prepared the figures; I.
Das, A. Leviton, and G. Zug read and commented on the manuscript.

LITERATURE CITED

ATHREYA, R. M., A. S. CAPTAIN, AND V. R. ATHREYA. 1997. A faunal survey of Namdapha Tiger Reserve,
Arunachal Pradesh, India. July, 1997. Unpublished report.

BISWAS, S. AND D. P. SANYAL. 1965. A new species of wolfsnake of the genus Lycodon Boie [Reptilia:
Serpentes: Colubridae] from the Andaman and Nicobar Islands. Proceedings of the Zoological Society, Cal-
cutta 18:137-141.

COLLINS, N. M., J. A. SAYER, AND T. C. WHITMORE. 1991. Atlas of tropical forests: Asia and the Pacific. Simon
and Schuster, New York. 256 pp.

DOWLING, H. G. AND J. V. JENNER. 1988. Snakes of Burma: Checklist of reported species & bibliography. Smith-
sonian Herpetological Information Service No. 76. 19 pp.

FITZINGER, L. J. F. J. 1826. Neue Classification de Reptilien nach ihren naturlichen Verwandtschaften. Wien.
66 pp.

LANZA, B. 1999. A new species of Lycodon from the Philippines, with a key to the genus (Reptilia Serpentes
Colubridae). Tropical Zoology 12:89-104.

LEVITON, A. E. 1965. Contributions to a review of Philippine snakes, VIII. The snakes of the genus Lycodon H.
Boie. Phillipine Journal of Science 94:117—140.

MANI, M. S. 1974. Ecology and biogeography in India. W. Junk, The Hague, Netherlands. 773 pp.

PAWAR, S. S. AND B. C. CHOUDHURY. 2000. An inventory of Chelonians from Mizoram, North-east India: New
records and some observations on threats. Hamadryad 25:144—158.

PAWAR, S. AND S. BISWAS. 2001. First record of the parachute gecko Ptychozoon lionotum Annandale 1905 from
the Indian mainland. Asiatic Herpetological Research (in press).

RAMAKRISHNAN, P. S. 1992. Shifting agriculture and sustainable development: an interdisciplinary study from
northeastern India, 1st ed. UNESCO, Paris.

RIPLEY, S. D. AND B. M. BEEHLER. 1990. Patterns of speciation in Indian birds. Journal of Biogeography
17:639-648.

SLOWINSKI, J. B. AND W. WUSTER. 2000. A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetologica
56:257-270.

SMITH, M. A. 1943. Fauna of British India, Ceylon, and Burma, including the whole of the Indo-Chinese
sub-region. Reptilia and Amphibia. Vol. III. Serpentes. Taylor and Francis, London. 583 pp.

STAMP, L. D. 1924. Notes on the vegetation of Burma. Geography Journal 64:23 1-237.

. 1930. Burma: an undeveloped monsoon country. Geographic Review 20:86—109.

TAYLOR, E. H. 1965. The serpents of Thailand and adjacent waters. University of Kansas Science Bulletin
45:609-1096.

TuN Nyo. 1997. Alaungdaw Katapha National Park (brief notes). Report to Nature and Wildlife Conservation
Division, Forest Department, Ministry of Forestry, Myanmar. 20 pp.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS
OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 21, pp. 407-509, 114 figs.

_ November tay 2001
Grenadiers (Families Bathygadidae and Macrouridae,
Gadiformes, Pisces) of New South Wales, Australia

By
Tomio Iwamoto
Department of Ichthyology, California Academy of Sciences
San Francisco, California 94118
and

Ken J. Graham
NSW Fisheries, P.O. Box 21, Cronulla, NSW 2230, Australia

[Warne Sctoueal Laboraton/

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 52, No. 21, pp. 407-509, 114 figs. November 13, 2001

Grenadiers (Families Bathygadidae and Macrouridae,
Gadiformes, Pisces) of New South Wales, Australia

by
Tomio Iwamoto

Department of Ichthyology, California Academy of Sciences,
San Francisco, California 94118

and

Ken J. Graham
NSW Fisheries, P.O. Box 21, Cronulla, NSW 2230, Australia

Twenty-six years of extensive trawling off New South Wales (NSW) by the FRV Kapala,
supplemented by opportunistic sampling by ORV Franklin, revealed a grenadier fauna
comprising 60 species in 21 genera. Despite a relatively limited coastline (600 nautical
miles), the diversity in NSW is comparable to those off Western Australia (63 spp.), New
Caledonia (63 spp.), and New Zealand (about 67 spp.). Of the 60 species, none are endemic
to NSW, but 12 are the only Australian records. Most of the material used in this study and
data on abundance, depth, and distribution were collected on the Kapala during fishery re-
source surveys. Between 1972 and 1997, Kapala surveyed shelf and slope depths along the
whole NSW coast, although most trawlirg was on the more extensive and commercially
productive grounds off central and southern NSW. The maximum depth trawled was
about 1200 m. Grenadiers were present in about 10% of tows on the outer-shelf (about
100-200 m), about half of all tows between 200 and 300 m, and in almost every trawl deeper
than 300 m. Smaller-meshed nets caught on average two more species per station than
those with larger mesh, and the mean number of species increased with depth. The data
also suggested that the NSW grenadier fauna is relatively rich in species at depths beyond
those sampled by Kapala. Depth-distribution data separated the species into two groups,
an upper-slope group in about 200—700 m, and a lower-slope group in 700-1300 m. Many
species that were rarely caught by Kapala appeared to be at the fringes of their geographic
or depth distributions in NSW. The large genus Caelorinchus was represent by 15 species,
while Coryphaenoides had 11 species. All other genera had four or fewer representatives.
Keys, figures, and brief accounts are provided for all NSW genera and species, with em-
phasis in the species accounts placed on their abundance and distribution in NSW.

Fishes of the families Bathygadidae and Macrouridae, often referred to as grenadiers, whiptails,
and rattails, are among the most abundant members of the demersal fish fauna at continental slope
depths of the world’s oceans. The rich grenadier fauna of the southwestern Pacific has only recently
come to the attention of the scientific world, mainly through the research and development of deep-
water trawl fisheries around Australia and New Zealand. McCann and McKnight (1980) made the
first major study of the New Zealand grenadier fauna and recognized 25 species from the area. That
number has been increased by the addition of new records and new species described by McMillan
and Paulin (1993)[Caelorinchus], McMillan (1995) [Trachyrincus], Iwamoto and McMillan (1997)
[ Trachonurus], and McMillan (1999) [Coryphaenoides]. Current research by McMillan and Iwamoto

407

408 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

suggests that the number will exceed 65. The New Caledonian grenadiers were treated by Iwamoto
and Merrett (1997) and Merrett and Iwamoto (2000). They recorded 63 species belonging to 20 gen-
era.

Australia’s rich fauna was documented little more than a decade ago by Paxton et al. (1989) as
having about 57 species of grenadiers. Unpublished records of Iwamoto suggest that there are more
than 100 species in all of Australia’s waters. Iwamoto and Williams (1999) found 63 species (17 new
species; 20 genera) off the western and northwestern coasts of Australia. On the southeast coast of
Australia, extensive deep-water trawling by the fisheries research vessel Kapala over a period of 26
years is principally responsible for revealing an extremely diverse grenadier fauna off New South
Wales (NSW). The number of species now known from NSW is 60 in 21 genera.

The diversity of the NSW fauna is unusually large given the limited extent of coastline [600 nauti-
cal miles (n. mi.)] and is probably attributable to the geographic position of NSW, between the spe-
cies-rich tropical waters to the north and the productive temperate waters to the south. The grenadier
fauna of NSW includes species known from the Coral and Tasman seas as well as from the Indian and
Southern oceans. Our knowledge of this diversity results from the extensive sampling by the Kapala
on the NSW slope over a prolonged period, which provided a unique opportunity for the collection of
grenadiers.

The purposes of this paper are threefold: (1) to record the species of grenadiers of New South
Wales; (2) to provide a key to the species and brief descriptions as an aid to their identification; and (3)
to record aspects of their distribution and abundance off NSW, so far as information is available.

Historical Perspective. Most of the grenadiers collected off NSW are housed in the Australian
Museum, Sydney (AMS). The earliest AMS specimens were collected in 1906 by the government
vessel Woy Woy from “a single cast of a small trawl which was built on the principal of one designed
and successfully used by the Prince of Monaco. It was lowered in 800 fathoms at a point thirty-five
miles due east of Sydney, on the 152nd Meridian” (McCulloch 1907). Grenadiers collected from this
trawl were Caelorinchus innotabilis (2 specimens), C. ‘fasciatus’ [maurofasciatus] (1),
“Macrourus”’ [Lucigadus] nigromaculatus (5), and “Optonurus” [Lepidorhynchus] denticulatus
(6). Caelorinchus innotabilis and Lucigadus nigromaculatus were subsequently described by
McCulloch as new species. It should be noted that the grenadiers and other teleosts collected with
them (see McCulloch 1907) are all upper-slope species, which suggests that the depth of this station
was, in fact, much less than the stated 800 fathoms and probably around 300 fathoms (550 m).

Between 1920 and 1968, a further 27 grenadier specimens were registered in the AMS collection;
these were C. australis (3), C. mirus (22), and L. denticulatus (2), all upper-slope species. The
holotype and four paratypes of C. mirus are included in this collection (see McCulloch 1926).

In 1970 the collection and study of NSW deep-water fishes was given great impetus with the
commissioning of FRV Kapala (Plate 1) by the NSW State Government. The 26 m Kapala was built
as a research vessel designed principally for trawling. Betweeen 1971 and 1997, Kapala conducted
numerous research surveys on continental shelf and slope trawling grounds between southern
Queensland and eastern Victoria. During this period, about 1260 grenadier specimens (55 spp.) from
217 Kapala stations in depths between 130 and 1240 m were registered in the AMS collection.

A further 40 grenadiers from off NSW were deposited in AMS by other collectors between 1970
and 1999; these included specimens of Coryphaenoides striaturus and C. filicauda from depths of
1590 and 2450 m. Additional NSW grenadiers are held by the Museum of Victoria (NMV) in Mel-
bourne (69 specimens, | 6 spp.); included are four species not represented in the AMS collections. The
I.S.R Munro Ichthyological Collection at the Commonwealth Scientific and Industrial Research Or-
ganization, Marine Research Laboratories, Hobart (CSIRO), also has a small collection of grenadiers
caught off NSW between 1975 and 1993 (41 specimens, | 1 spp.), but this collection contains no addi-
tional species to those in AMS and NMV.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 409

PLATE 1. The 26-meter fishery research vessel Kapala.

This paper describes the NSW grenadier fauna as represented in the AMS and NMV collections.
As the great majority of NSW grenadiers were collected from Kapala stations, much of the discussion
relates to observations made during Kapala’s offshore surveys. Relative abundance, depth, and distri-
bution data were collected by Graham on Kapala between 1972 and 1997, and during subsequent
trawling on a commercial trawler off the southern NSW port of Bermagui in 1999-2001.

MATERIALS AND METHODS

Study Area. New South Wales is on the east coast of Australia between latitudes 28°10’S and
37°45'S (Fig. 1). New South Wales waters extend to the edge of the Australian Fishing Zone (AFZ;
200 n. mi. from land) and include the area around Lord Howe Island (31°30’S, 159°05’E). Apart from
a few isolated seamounts and reefs, most of the seabed beyond the continental slope is deeper than
2000 m; some parts of the Lord Howe Rise within the AFZ are as shallow as 1000 m. The NSW conti-
nental shelf is narrow, with the shelf break mostly between 15 and 25 n. mi. offshore at a depth of
about 200 m. The continental slope (between the 200 and 2000 m isobaths) can be arbitrarily divided
into three depth zones. The upper-slope (200-700 m) is mostly between three and five n. mi. in width,
with a gradient between 1:10 and 1:20, while the mid-slope (700-1300 m) and lower slope (>1300 m)
are relatively much narrower and steeper.

The Kapala collected grenadiers from shelf and slope waters between 27°50'S and 38° 15'S over
a depth range of 130 to 1240 m. Collections were also made from a few mid-slope and lower-slope sta-
tions between 800 and 2500 m off Nowra by CSIRO’s oceanographic research vessel Franklin (NMV
specimens). A small number of grenadiers at AMS were collected by Franklin with a beam trawl and
epibenthic sled on the Lord Howe Rise.

410

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

ic
Queensland Danger PY)
ER see ey ie ee ee I CS i 29°
Clarence River] ‘Y

pa
ia

LW)
31°

boy Crowdy Heads
SOUTH Tuncurry
WALES Port Stephens ue

Newcastle Aa 4 ea

te
Broken Bay Ze

Sydney = oo Tasman

Wollongong We Sea
i af
i a

Nowra =
= 35°

Jewis Bay
Ulladulla f

Batemans Bay ce
_ }eMontague I.
Bermagui i

5. eden

Nic. Gabo |

152° 154°

150°

FIGURE |. Map of New South Wales coast showing FRV Kapala stations where grenadiers were collected for AMS.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 411

Kapala Trawl Gear and Survey Methods. A range of net sizes and styles was used during the
many Kapala exploratory and stock-assessment surveys, with the choice dependent on the purpose of
each study. Nets for the capture of prawns ranged in size between 20 and 30 m headline length and
were constructed throughout with 45 mm mesh netting. Fish trawls had headline lengths between 20
and 56 m, and were made with relatively large-mesh panels in the front of the net (100-200 mm mesh)
and usually with codends of 90 mm mesh. During some surveys, fish trawls were fitted with 45 mm
mesh codend liners. Towing speed in upper-slope depths was between 2.5 and 3.5 knots; mid-slope
trawling was usually 1.8—2.5 knots. The maximum depth trawled by Kapala (1240 m) was limited by
the amount of trawl warp carried on her main winch (2500 m). A large mid-water trawl was also de-
ployed in oceanic waters off Sydney-Newcastle on a few occasions in 1977-79 for the capture of
bathypelagic species.

Normal practice for exploratory and stock-assessment trawling was to tow along a selected depth
for one to two hours. The depth range of upper-slope tows was usually within + 20 m of the target
depth, while on the steeper mid-slope the range was often greater, around + 40 m of the target. Re-
corded fishing depths were for the period the trawl was fishing the seabed. Catches were sorted into
commercial and non-commercial species for assessment; all grenadiers caught off NSW were consid-
ered to be part of the non-commercial component of the catch. A list of all fishes (with approximate
numbers) was compiled for each station. Any rarely caught specimens were retained for AMS.

TAXONOMIC DESCRIPTIONS

In the descriptive section of this work, general features of the family, subfamily, and genera are
provided separately under each section and in the key. Detailed characters that distinguish the species
are given under the species descriptions, although characters previously given in the keys are gener-
ally not repeated except for necessary elaboration. For synonymies and additional figures and descrip-
tions of most of the species and genera treated here, the reader is referred to Last et al. (1983), Gomon
et al. (1994), Iwamoto and Merrett (1997), Iwamoto and Williams (1999), Merrett and Iwamoto
(2000), and references cited in the last three publications. Methods of taking measurements and
counts are described in detail in Iwamoto and Sazonov (1988) and in a condensed version in Iwamoto
and Williams (1999).

Abbreviations for fins are 1D = first dorsal, 2D = second dorsal, P = pectoral, V = pelvic. Spinous
rays in the first dorsal fin are designated with Roman numerals, segmented rays are given in Arabic
numerals (e.g., II,10). The splintlike uppermost ray of the pectoral fin is designated with a small i(e.g.,
il6). Counts of gill rakers (GR) are distinguished as to which gill arch the rakers are counted (e.g., first
or outermost arch = GR-I, second arch = GR-II) and whether the outer or inner series is counted.
Gill-raker counts are usually made on the right side of the fish, with the gill covers pulled back to ex-
pose the rakers, which are often quite small. In some species, the upper connection of the gill cover to
the body must to severed in order to pull the cover back far enough to expose the rakers. Scale rows are
counted in a diagonal series from the first dorsal origin (“below 1D’), mid-base of 1D, and below 2D
origin, to, but not including, the lateral-line scale. The count of lateral-line scales includes those from
the anterior origin of the lateral line to a point marking the distance equal to that from the snout tip to
the origin of the first dorsal fin (viz., the predorsal length) (Fig. 2). Size given is maximum total length,
rounded up to the nearest 5 cm. Institutional abbreviations follow Leviton et al. (1985) and Leviton
and Gibbs (1988). Australian states are abbreviated as follows: New South Wales, NSW; Queensland,
Qld; South Australia, SA; Tasmania, Tas.; Victoria, Vic.; Western Australia, WA.

412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

scale rows below 1D
predorsal length :
scale rows below mid-base 1D

scale rows below 2D

preoral
length

lateral line scales
over distance equal to predorsal length

FIGURE 2. Diagrammatic illustration of typical grenadier to show method of counting scale rows and measuring predorsal
length, preoral length, and suborbital width; (a) lateral view; (b) dorsal view of head. Abbreviations: in, internasal width, io,
interorbital width, po, postorbital length, op, opercle to preopercle distance, 1D, first dorsal fin, 2D, second dorsal fin.

RESULTS

Kapala Surveys. Prior to 1970, little was known about the extent and productivity of trawl
grounds along the NSW slope. Between 1971 and 1981 exploratory trawling and resource assessment
surveys determined the extent of upper-slope trawling grounds and the nature of demersal prawn and
fish stocks on those grounds (Gorman and Graham 1975; Graham and Gorman 1985; Andrew et al.
1997). Off northern NSW only two relatively small areas of trawlable seabed were found, one off the
Clarence River and the other off the NSW-Queensland border. Apart from a single tow in 740 m off
Danger Point, all trawls north of latitude 31°30’S were shallower than 600 m. Because catch rates of
commercial fishes were relatively low during early surveys, there was no trawling by Kapa/a in any
slope depths offnorthern NSW after 1978. At present, commercial trawlers occasionally fish for royal
red prawns (Haliporoides sibogae) and deepwater slipper lobsters (Jbacus spp.) in 200-500 m on the
Clarence River ground.

In contrast, relatively large areas of trawlable ground with commercial fish stocks were found on
the upper-slope off central and southern NSW. A significant trawl fishery for both deep-water fishes
and prawns developed during the late 1970s (Tilzey 1994), and today about 40 trawlers continue to
fish these grounds. Consequently the majority of Kapala upper-slope trawling was done south of Port
Stephens, particularly on grounds off Sydney, Ulladulla, and Eden-Gabo Island. This included the fi-
nal study by Kapala before her decommissioning in 1996-97, which was a repeat stock-assessment
survey of the upper-slope south of Newcastle (Graham et al. 1997, 2001).

Mid-slope trawl grounds (700-1200 m) were charted in detail by Kapala between 1983 and 1989
(Graham 1990). Trawlable seabed on the mid-slope was more restricted in area than on the up-
per-slope, and very little trawl ground was found in depths below 1000 m. Most fishable ground was
located between latitudes 31°40'S and 36°00’S, and consequently, almost all mid-slope stations were
in this area. A small number of tows were also made southeast of Gabo Island (37°40'S). No large

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 413

stocks of commercial fish were found, but commercial trawlers occasionally target orange roughy
(Hoplostethus atlanticus Collett, 1889), oreo dories (family Oreosomatidae), and edible dogsharks
(family Squalidae) in mid-slope depths off NSW.

Outer shelf grounds (100-200 m) were surveyed off northern NSW in 1978, and between Port
Stephens and Gabo Island in 1993-94 (Graham et al. 1995, 1996).

Kapala Grenadier Data. Table | shows the distribution by depth and latitude of Kapala survey
trawls on the NSW outer shelf and slope between 1976 and 1997. For each of these trawls a list of
fishes was compiled, and from these data, abundance and distributional information for NSW grena-
diers were derived (see species descriptions for details). Figure | shows the locations of all Kapala
stations from which grenadiers were collected for AMS; station numbers, location and depths for
these are listed in Appendix 1.

The depth and geographic ranges derived from Kapala catches were influenced by the distribu-
tion of stations, which, as discussed above, reflected the lack of mid-slope trawling off northern NSW.
As Table | shows, most trawling was done on the commercially productive grounds off central and
southern NSW, the maximum depth trawled was about 1200 m, and no depths greater than 740 m
were sampled north of Crowdy Head (31°45’S). The style of trawl used during the various surveys
may have also influenced the data. About 75% of upper-slope trawls and 56% of mid-slope trawls
were done with large-meshed fish nets, which reduced the likely capture of small specimens. The cap-
ture rate by Kapala may therefore understate the true abundance of some species of small adult size.

During the period that detailed catch data were recorded (1976-97), grenadiers were caught in
1072 Kapala trawls over a depth range of 130-1240 m. Grenadiers were present in about 10% of
outer-shelf tows, about half of all tows between 200 and 300 m, and in almost every trawl deeper than
300 m. The number of species caught at each station was related to the trawl gear and depth fished
(Fig. 3). Nets with 45 mm codend mesh caught, on average, two more species per station than those
with 90 mm mesh, and the mean number of species per trawl increased almost linearly with depth (see
Appendix 2 for data). The data also suggest that the grenadier fauna off NSW is relatively rich in spe-

TABLE |. Distribution by latitude and depth of FRV Kapala demersal stations over 100 m trawled between
1976 and 1997. New South Wales distribution and abundance were derived from catches at these stations.

Depth (m)

Latitude 100—- 200—- 300—- 400— 500— 600— 700— 800— 900— 1000— 1100— Total
(°S) SE = 2OSRI S99 Fe TAO O SOD 699M FOO! 8995 "999%" 51099. 1200

27 0 | | 0 l 0 0 0 0 0 0 3
28 22 3 3 5 2 0 l 0 0 0 0 36
Zz 14 a 9 13 A 0 0 0 0 0 0 47
30 11 7 l 0 0 0 0 0 0 0 0 19
31 6 2 l 0 0 0 0 l l l 0 2
32 94 8 10 13 9 l 4 10 14 3] 0 194
33 104 45 68 96 34 14 15 22 30 23 6 457
34 102 10 13 44 10 i | 8 18 12 4 235
35 79 53 64 54 33 10 4 ) 24 Is 3 348
36 71 9 9 6 0 0 0 0 0 0 0 95
a7 iS 42 68 45 22 6 l 3 3 2 0 267
38 | 4 10 10 6 3 l 0 ] 1 0 37

ome |
°
-
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414 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Mean No. of Species

eee ape peepee 2S
ae
—e 45 mm codend
—l— 90 mm codend

150- 250 350 450 550 650 750 850) 950\0 M@aQiaines
Depth (m)

=n
i)

Oo

Mean No.
[@>)

FiGuRE 3. Graph showing influence of depth and mesh size to mean number of grenadier species captured off New South
Wales by Kapala.

cies at depths beyond those sampled by Kapala. Apart from a small number of ORV Franklin stations,
the lower slope and extensive abyssal seabed off NSW is largely unsampled.

Figures 4 and 5 summarize the depth and geographical ranges for each of the 55 species caught by
Kapala. In Figure 4, the 53 demersal species are ordered according to their minimum depth of capture;
those recorded on more than five occasions are divided into “upper-slope” and “mid-slope” groups,
and rarely caught species are listed beneath. The same species, ordered by their latitudinal ranges, are
shown in Figure 5; the bathypelagic species Cynomacrurus piriei and Odontomacrurus murrayi are
included in this figure. The depth range of the upper slope is arbitrarily defined as 200—700 m and the
mid slope as 700-1300 m. Figure 4 shows that while some species fit neatly within these depth cate-
gories, many show some overlap, and for those taken in the deepest trawls (>1100 m), no maximum
depth range can be defined. There is an overall pattern of gradually increasing depth ranges, but within
this a number of depth groupings can be discerned.

Ten species are characteristic of the upper slope. Although the depth ranges of six of these ex-
tended to about 800 m or more, each was most abundant in depths less than 700 m. The two species
with the shallowest depth ranges, Caelorinchus mirus and C. australis, were also caught in outer-shelf
depths; either or both species were present in about 10% of the 579 outer-shelf trawls.
Lepidorhynchus denticulatus was the most abundant upper-slope species and also exhibited the great-
est depth range (230-1080 m) of any NSW grenadier. Trends in geographic range are also evident for
several upper-slope species (Fig. 4). The ranges of three species (Lucigadus microlepis, Ventrifossa
nigrodorsalis, and Hymenocephalus longibarbis) with known tropical distributions extended to cen-
tral or southern NSW; the latter two were caught by Kapala as far south as Jervis Bay and Ulladulla,

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 415

No. of Depth (m)
Records 100 200 300 400 500 600 700 800 900 1000 1100 1200

ee ee

Upper-slope Species

Caelorinchus australis 77 ———
Caelorinchus mirus 293 ——ee
Caelorinchus parvifasciatus 239 eS
Lepidorhynchus denticulatus 601 —_—_—_—_—$_=—_—_—_—_—_——o_—_————
Caelorinchus maurofasciatus 254 —_—_—_—_—_—_—_—_—_—_—_=
Malacocephalus laevis 289 [SSS
Ventrifossa nigrodorsalis 68 EES
Hymenocephalus longibarbis 102 —_—
Lucigadus nigromaculatus 222 —_—_—_—_—_—_—_—_—_—_—_—_—_—_——————
Lucigadus microlepis 14 ———
Mid-slope Species
Caelorinchus innotabilis 227 ————
Caelorinchus macrorhynchus 9 —_——_—_——_—_—_—__
Caelorinchus fasciatus 8 Se
Nezumia propinqua 43 —_—_————
Kuronezumia bubonis 20 —_—_—_—_——__
Ventrifossa johnboborum 23 ee
Gadomus sp. cf. colletti 28 ——
Caelorinchus matamuus 41 ———
Coryphaenoides dossenus 185 Se
Coryphaenoides serrulatus 253 —_—_—_—_—_—_—__—_—
Coryphaenoides subserrulatus 173 nd
Mesobius antipodum 168 eS
Nezumia namatahi &/or N. kapala 133 i
Nezumia namatahi (AMS Records) 9 $$
Nezumia kapala (AMS Records) 15 ——
Kuronezumia leonis 146 —_—_—_—_—_—_—_—_—__
Caelorinchus acanthiger 224 ee ———————
Ventrifossa paxtoni 29 —SSaees
Gadomus pepperi 178 ee
Nezumia coheni 57 SSS SSS
Caelorinchus kaiyomaru 148 en
Trachonurus gagates 63 es
Sphagemacrurus richardi 17 ———se
Bathygadus cottoides 26 en
Cetonurus globiceps 34 en
Haplomacrourus nudirostris 8 ——==
Bathygadus furvescens 11 ——
Rarely Caught Species
Caelorinchus sp. cf. cingulatus 4 —
Caelorinchus supernasutus 2 —
Caelorinchus smithi 1 —
Caelorinchus kermadecus 5 Se
Mataeocephalus spp. 3 a
Hymenocephalus aterrimus 2 rs
Hymenocephalus nascens 2 SS
Trachonurus sentipellis 4) re
Caelorinchus mycterismus 2 —=
Coryphaenoides filicauda 1 -
Coryphaenoides grahami 4 er
Coryphaenoides rudis 2 ——
Trachyrinchus longirostris 4 ———
Coryphaenoides striaturus 3 —
Bathygadus sp. cf. spongiceps Z =sssnss
Caelorinchus trachycarus 1 =
ee ee ee | ee es ee

100 200 300 400 500 600 700 800 900 1000 1100 1200

Ficure 4. Depth distributions for NSW species collected by Kapala (thin line represents range or range extension by | or 2
observations only).

416 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

No. of Latitude (°S)

Records 280 = 2906 sO ol 32 33) 8S4 OC OMS aS
Uppersiope Species ==S—S=*”*=CStsi‘“‘i‘“S~‘~C SC ee
Caelorinchus smithi 1

Caelorinchus sp. cf. cingulatus 3 —=
Lucigadus microlepis 14 —
Ventrifossa nigrodorsalis 71 ee
Hymenocephalus longibarbis 102 a
Malacocephalus laevis 290 ——
Caelorinchus mirus 293 (EET
Lucigadus nigromaculatus 222 a ,
Lepidorhynchus denticulatus 603 EES
Caelorinchus parvifasciatus 239 SS
Caelorinchus maurofasciatus 254 ee
Caelorinchus australis 77 —
Mid-slope Species
Coryphaenoides filicauda 1 -
Hymenocephalus aterrimus 2 =
Hymenocephalus nascens 2 —
Mataeocephalus spp. 3 —
Trachonurus sentipellis 2 —
Sphagemacrurus richardi 18 a
Caelorinchus mycterismus 2 rs
Bathygadus furvescens 11 et
Caelorinchus macrorhynchus 8 ee
Caelorinchus supernasutus 2 nt
Haplomacrurus nudirostris 8 eee
Kuronezumia bubonis 23 ee
Ventrifossa johnboborum 23 =e ES
Bathygadus cottoides 26 eS
Caelorinchus kermadecus 5 =
Cetonurus globiceps 34 pee
Coryphaenoides grahami 4 a
Gadomus sp. cf. colletti 27 ee
Nezumia propinqua 43 es
Ventrifossa paxtoni 29 ape ee
Caelorinchus acanthiger 224 ee
Caelorinchus innotabilis 227 eS
Caelorinchus kaiyomaru 148 SE SES
Coryphaenoides dossenus 143 —_—_—_—_—_—_—_—_—_—_—_———
Coryphaenoides serrulatus 253 SS nS
Coryphaenoides subserrulatus 173 aS
Gadomus pepperi 178 a
Kuronezumia leonis 147 en
Mesobius antipodum 168 ee
Nezumia coheni 60 eS
Nezumia kapala &/or N. namatahi 133 ES
Nezumia kapala (AMS Records) 15 (ee
Nezumia namatahi (AMS Records 9 ————
Trachonurus gagates 65 —————— ey
Coryphaenoides rudis 2 —
Trachyrinchus longirostris 4 —
Caelorinchus matamuus 44 ——————— ae
Bathygadus sp. cf. spongiceps 2 —=
Caelorinchus fasciatus 8 ———
Coryphaenoides striaturus 3 =
Caelorinchus trachycarus 1 -
Bathypelagic Species
Odontomacrurus murrayi 4 ee
Cynomacrurus piriei 1 =

es es See) ee Ce eee Saeeen Seeeee) eee! 2

28 29 30 31 32 33 34 35

Ww
(o>)
WO
N
Ww
foe}

FIGURE 5. Geographical distributions for NSW species collected by Kapala.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 417

although recently both were collected (by Graham) further south off Bermagui (36°20'S). Four south-
ern species showed limits to their northern distributions within NSW: C. parvifasciatus and
Lepidorhynchus denticulatus did not extend north of the Clarence River, C. maurofasciatus was only
caught south of Port Stephens, whereas C. australis was restricted to waters south of Batemans Bay.

The depth range of C. innotabilis (450—1075 m) spanned both the upper and middle slopes, but
the species was most abundant between 600 and 900 m. Caelorinchus innotabilis can be included in a
group that mainly inhabits the shallower mid-slope depths. This group also includes Caelorinchus
macrorhynchus, C. fasciatus, C. matamuus, Gadomus sp. cf. colletti, Kuronezumia bubonis, Nezumia
propinqua, and Ventrifossa johnboborum. They were mostly caught shallower than about 1000 mand
mainly between 700 and 900 m, and although all showed relatively extensive geographic ranges, none
was abundant in any part of their NSW range. Four of these species are also found in tropical waters; of
these, C. macrorhynchus, K. bubonis, and V. johnboborum were caught by Kapala south to Jervis
Bay, and N. propinqua to about Batemans Bay. A juvenile C. macrorhynchus was recently collected
(by Graham) further south near Montague Island. Two southern species, C. fasciatus and C.
matamuus, were taken as far north as Broken Bay.

Eighteen species (with eight or more captures) were found only in depths greater than 700 m.
Haplomacrourus nudirostris, Sphagemacrurus richardi, and Ventrifossa paxtoni are species previ-
ously reported from more tropical waters than NSW (Iwamoto and Merrett 1997) and were caught by
Kapala mostly north of Sydney and only between 800 and 1100 m. The full depth ranges of the re-
maining species were probably greater than those shown by Kapala catches. Six species,
Coryphaenoides dossenus, C. serrulatus, C. subserrulatus, Kuronezumia leonis, Mesobius anti-
podum, and Nezumia namatahi, were caught across the full mid-slope depth range (about
700-1200 m) and on all grounds between Crowdy Head and Gabo Island. Another five species also
caught between Crowdy Head and Gabo Island but with greater minimum depths (800—900 m) were
Caelorinchus acanthiger, C. kaivomaru, Gadomus pepperi, Nezumia coheni, and Trachonurus
gagates; N. kapala is also likely to be in this group (based on AMS collection; see species descrip-
tion). These 12 species have generally southern distributions, being found around the south coast of
Australia and many also off New Zealand. The last three mid-slope species on the list, Bathygadus
cottoides, B. furvescens, and Cetonurus globiceps, were mostly caught deeper than 1000 m and are
possibly more abundant at depths greater than trawled by Kapala.

Species rarely caught by Kapala may be put in one or other of the groups discussed above, but
there are too few observations to be definitive. Most of these species appear to have been caught at the
fringe of either their geographic or depth range. Eight species, Caelorinchus cingulatus, C.
kermadecus, C. smithi, Hymenocephalus aterrimus, H. nascens, Mataeocephalus sp., M.
acipenserinus, and Trachonurus sentipellis, are more commonly found in tropical waters (Iwamoto
and Merrett 1997). Kapala captures of these species were from the most northern stations, although
the small size of Hymenocephalus spp. and Mataeocephalus spp. may have also contributed to their
relatively low number of captures. In contrast, Cynomacrurus piriei, which was caught once by
Kapala off the NSW south coast, is primarily a Southern Ocean species (Iwamoto 1990). McMillan
and Paulin (1993) reported that Caelorinchus mycterismus and C. supernasutus are relatively com-
mon around northern New Zealand and are also recorded from the Wanganella Bank to the northwest
of the North Island. The Kapala captures are the only confirmed specimens from the western Tasman
Sea. The few specimens of Bathygadus sp. cf. spongiceps, Caelorinchus trachycarus,
Coryphaenoides filicauda, C. grahami, C. rudis, C. striaturus, and Trachyrincus longirostris were
caught only in trawls deeper than 1000 m and are possibly more abundant at depths greater than those
sampled by Kapala.

418 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

TAXONOMY

We use grenadier as a collective term for four distinct groups of gadiform fishes, each of which
has at one time or another been considered as a separate family. In fact, the bathygadids have been
placed in a separate suborder (Gadoidea) from the others (Macrouroidea) (Howes 1989). The prob-
lems of relationships have yet to be adequately resolved, so for this paper we have chosen to take a
conservative approach as used by Iwamoto and Merrett (1997). It should be noted that the long-tailed
hakes (Macrouronus spp., Merlucciidae) are also called grenadiers. The southern Australian species,
M. novaezealandia, is known as blue grenadier (Last et al. 1983; Gomon et al. 1994). All grenadiers
have a long, tapered tail with long dorsal and anal fins that meet posteriorly without a noticeable cau-
dal fin, and the anal fin lacks an elevated lobe. The trachyrincines are reported to have a caudal fin, but
it is so rudimentary that examination under magnification is usually necessary to observe its presence.
One often finds specimens with what appears to be a sizable caudal fin, but that structure is a result of
loss of the tail tip and an overgrowth of the anal and dorsal fin rays. Grenadiers can be distinguished
from other deep-sea fishes that have a long tapered tail by a combination of the lack ofa distinct caudal
fin, one or two dorsal fins, a single, long, low anal fin lacking lobes or dips in its profile, pelvic fins
present (in all but Macrouroides inflaticeps), well separated, and consisting of five to as many as 18
rays, and no teeth on roof of mouth or on tongue.

KEY TO THE FAMILIES AND SUBFAMILIES OF GRENADIERS FROM NEW SOUTH WALES

la. A single long-based, low dorsal fin; head enormous, bulbous and spongy; eyes tiny, more than 10 in head length
(E1G56) ores, teen eet cee [Macrouridae, subfam. Macrouroidinae—not yet found off NSW but can be expected]
1b. Two dorsal fins; head variously shaped, not especially huge and bulbous; eyes less than 10 in head length. ...... 2

2a. First and second dorsal fins separated by a distinct gap (Fig. 7a); gill rakers all short, tubercular or tablike (Fig. 8a);
opening of first gill slit restricted by membrane across upper and lower arms of gill arch (Fig. 9)
Siecle o et Jcitdaste Ome innate cee te ene he Re ear ee Macrouridae, subfam. Macrourinae (Fig. 10)
2b. First and second dorsal fins closely approximated, without a distinct gap (Fig. 7b); gill rakers slender, lathlike
(E1g58b)iopeninpiofefirst pillishtiunrestricted’by memibrane! | © -)as- es ee ee eee 3

3a. Mouth subterminal to inferior, a long, stout, sharp snout; scales covered with spinules; series of sharply spined
scutes present along bases of dorsal and anal fins (Fig. 11)............ Macrouridae, subfam. Trachyrincinae
3b. Mouth essentially terminal, snout rounded not protruding (Fig. 12); scales all smooth; no scutes on body. Bathygadidae

F\GURE 7.(a) Firstand second dorsal fins separated by a distinct gap; (b) first and second dorsal fins closely approximated.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 419

tubercular
gill rakers sect
AX. € 5 ke
oe v ws pecs gill rakers
ler \
] \\\\ \\ Wy
WLLL ;

FiGure 8. (a) Gill rakers on outer arch short and tubercular; (b) gill rakers long and slender.

outer gill slit

Ficure 12. Diagrammatic illustration of a Bathygadidae (Bathygadus sp.).

420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

FAMILY BATHYGADIDAE

DISTINGUISHING FEATURES. — Two dorsal fins, the second beginning immediately behind first
without a pronounced gap. Outer gill rakers on first arch long, lathlike, 20 or more total.
Branchiostegal rays 7. No membrane restricting first gill slit. Pelvic fin rays 8-10; first dorsal fin with
smooth, flexible spinous ray. No spinules on scales. Large terminal mouth and no protruding snout.

REMARKS. — Only two genera are recognized, Bathygadus and Gadomus, with about 35 nomi-
nal species; the taxonomy of the group is still not fully resolved. Members of the family are found in
tropical to temperate waters, and the family is widely distributed in the Atlantic, Indian, and Pacific
oceans, but notably absent (as are many macrourid genera) in the eastern North Pacific. Bathygadids
are abundant throughout the Indian Ocean and the western South Pacific, but their occurrence and
abundance in the central and eastern South Pacific is uncertain because of the lack of sampling in those
regions (aside from the Nazca and Sala y Gomez ridges in the southeastern Pacific; see Sazonov and
Iwamoto 1992). Contrary to Howes and Crimmen’s (1990:201) statement, the family is not found in
the Southern Ocean as it is usually defined (see for example, Gon and Heemstra 1990), as the
Kermadec Islands and Lord Howe Rise, from which they cite occurrences of the family, have never
been considered part of the Southern Ocean.

REFERENCES. — Gilbert and Hubbs (1920); Howes and Crimmen (1990); Iwamoto (1990).

KEY TO GENERA AND SPECIES OF BATHYGADIDS FROM NEW SOUTH WALES

la. Chin barbel well developed; ground color of body cream to light brown; first dorsal, pectoral, and pelvic fins with

stout\ greatly producedirays|(Fig! 13) i-wia ci oes Go a hs ce eee (Gadomus) 2
1b. Chin barbel absent; ground color of body dark, usually blackish; rays of first dorsal, pectoral, and pelvic fins weak,
anterior ray of fins if produced, distally hair thin(Fig.14)........................ (Bathygadus) 3
2a. Outer gill rakers 24-25 total, longest about length of longest gill filaments; chin barbel about 3 times orbit diameter;
mouth pale; all fins black; pyloric caeca short, more than100.................. Gadomus sp. cf. colletti
2b. Outer gill rakers 28-31 total, longest about twice length of gill filaments; chin barbel about 1—2 times orbit diameter;
mouth black; fins dark but not intensely black; pyloric caeca long, about75 ............. Gadomus pepperi
3a. Head broad, interorbital width 30-40% (Fig. 15a); orbit diameter 16-22%; paired fins dusky to black; flesh soft, head
bonesiweakeeee, .\si.c.rp Sekt Rete ees Sed SG ees see dae 2.5.8 @ a, a aoe ee 4
3b. Head narrow, compressed, interorbital width 26-30% (Fig. 15b); orbit diameter 21—23%; paired fins black; flesh firm,
headibones:ratherstrong). 2.5 25. 3 Shoat Se no ee ee B. furvescens
4a. Pyloric caeca 8-12; P 110-115 (usually 112-114); length longest gill rakers 14-20% HL; suborbital width 16-19% HL;
smaltfadultsizes<30%CMP ra costs etka, ec Sn Bo ee ae besa Re. Ae B. cottoides
4b. Pyloric caeca 15—28, usually 20-28; P 114-118; length longest gill rakers 7-16% HL; suborbital width 12-16% HL;
largeradultsize, >4Gcmi . 2. a 2 De eek bo ee ee 2 B. sp. cf. spongiceps

FiGURE 13. Diagrammatic illustration of a Gadomus sp. FIGURE 14. Diagrammatic illustration of a Bathygadus sp.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 42]

Genus Bathygadus

DISTINGUISHING FEATURES. — Distin-
guished from Gadomus by absence of chin bar-
bel, more fragile head bones and head covering,
generally darker color (usually mostly
blackish), absence of extremely long,
well-developed rays in dorsal and pectoral fins.

REMARKS. — Three species known from
NSW, each widely distributed in parts of the In-
dian and Pacific oceans; two species found in
other parts of Australia.

REFERENCES. — Gilbert and Hubbs
(1920); Howes and Crimmen (1990).

Bathygadus cottoides Ginther, 1878
Fig. 16

DISTINGUISHING FEATURES. — D II,8—10,
Pil0-114, rarely i15, V9 (rarely 8 or 10); outer — Figure 15. Comparison of internasal width (IN) and
GR-I (4-6)+(19-21), total 25-28, GR-II interorbital space (IO) in (a) Bathygadus furvescens and (b)
(outer) 17-20 total; pyloric caeca 8-12. Mea- 2athygadus sp. cf. spongiceps.
surements in percent HL: snout length 30—36;
internasal width 3 1—35; interorbital width 3 1-40; orbit diameter 16—21; suborbital width 16—19; dis-
tance orbit to angle of preopercle about 49-59 (sometimes less); upper jaw length 49-59; height as-
cending premaxillary process |3—16; vent to anal fin origin 9-29; length pectoral fin 41—86; length
pelvic fin 49-66; length outer gill raker 14—20. Outer pelvic ray in some specimens moderately pro-
longed, but most others lack produced rays; pectoral fin rays not extending beyond anus. Teeth bands
relatively narrow, 7 or 8 teeth wide at broadest part of premaxillary band, 4 or 5 wide in broadest part
of dentary band. Fin rays dark dusky to black. Flesh and head bones rather soft and weak.

SIZE. — Maximum size about 30 cm.

DISTRIBUTION. — Australia (NSW, Vic., Tas., SA, WA), New Zealand and southern Africa, in
depths of about 1000 m to more than 1500 m.

NSW CAPTURES. — Taken by Kapala between Crowdy Head and Batemans Bay in depths
greater than 950 m. One specimen (AMS 1I.29318-001) captured in 1325 m by ORV Franklin near
Lord Howe Island. Kapala captured 56 specimens in 26 tows (or 14% of Kapala tows deeper than
900 m). Because of its small adult size (most specimens less than 20 cm), B. cottoides is probably
more abundant than its trawl capture rate suggests.

REMARKS. — This species can be confused with small individuals of B. furvescens, but the lower
counts of pectoral fin rays and pyloric caeca, the wider head, and the smaller orbit of B. cottoides are
characters that differentiate the two species.

REFERENCE SPECIMENS. — AMS I.24978-005 (3 spec.); K84-20-04. AMS 1.25095-004 (1
spec.); K84-20-05. AMS 1.26000-010 (1 spec.); K86-01-08. AMS I.27638-003 (1 spec.); K88-11-01.
AMS 1I.27717-001 (1 spec.); K88-04-08. AMS 1I.29310-001 (1 spec.); ORV Franklin 28°44.08’'S,
161°54.59’E; 1325 m; 4 May 1989. AMS 1.29745-007 (1 spec.); K89-18-02. AMS I.29801-002 (1
spec.); K89-08-02. AMS 1I.39052-002 (3 juveniles); K89-18-04.

REFERENCES. — Iwamoto and Merrett (1997)(in part); Merrett and Iwamoto (2000).

422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

FiGurE 16. Bathygadus cottoides Ginther, 1878. AMS 1.24978-005. From Kapala stn K84-20-04, off Broken Bay, NSW, in
1070-1125 m.

Bathygadus furvescens Alcock, 1894
Fig. 17

DISTINGUISHING FEATURES. — Holotype data in square brackets [ ]. D II,8—9 [10], P 115-19
[il6], V 8-9 [8]; GR-I (outer/inner) (5—6)+(18—20)[6+18]/(3—4)+(15—16)[3+16], totals 23—26/
18-19, GR-IT 2+(14—15)[2+15 ]/(2—3)+(14—-16)[2+15], totals 16—17/17—18; pyloric caeca 20—22[20].
Measurements (in percent HL): snout length [26] 28-30; internasal width 24; interorbital width [26]
29-30; orbit diameter [21] 21—23; suborbital width 14—15; postorbital length [51] 51—52; distance or-
bit to preopercle [48] 48-50; length upper jaw [57] 56—59; length pectoral fin [61] 71-81; length pel-
vic fin 67-83; length longest gill raker 14-15. Fins well developed; pectoral and pelvic fins long,
extending to or beyond origin of anal fin; pelvic fin with distally filamentous elongated outer ray.
Flesh and head bones relatively stout and more like that of Gadomus. Teeth tiny, in broad villiform
band in both jaws, premaxillary band about 10 to 12 teeth at widest point, dentary band about 7 or 8
teeth at widest. Paired fins black, median fins black to dark dusky.

SIZE. — To approximately 55 cm.

DISTRIBUTION. — Known only from the holotype taken off the Maldives in 1315 m and the cur-
rent specimens from NSW and Tasman Sea, but can be expected in other parts of the Indian Ocean and
western Pacific. In addition to Kapala material, AMS specimen (1.29338-002) was captured in
1050 m on the Lord Howe Rise to the east of the Australian Fishing Zone.

NSW CAPTURES. — Kapala caught 14 specimens of B. furvescens at 11 stations between
Crowdy Head and Jervis Bay, in depths between 1000 and 1240 m. It was present in only 11% of all
Kapala tows deeper than 1000 m, but is possibly more abundant at greater depths.

REMARKS. — These NSW specimen agree rather closely with the holotype, for which count and
measurement data in the Distinguishing Features section were provided by Yuri I. Sazonov
(ZMMGU) and Yuri N. Shcherbachev (IOAN). The snout length, interorbital width, and pectoral fin
length in the holotype were slightly shorter than in the NSW specimens examined, and the count of
first dorsal fin rays was high. Alcock (1894:14) gave the pyloric caeca count as 20 (Sazonov and
Shcherbachev counted only 18). These data fall well within the expected range of variation. The pec-
toral fin length in the holotype may have been longer in the fresh specimen than when examined by
Sazonov and Shcherbachev. Alcock (ibid. ) stated that the fin“... tips reach beyond the origin of the
anal” and “the length . . . is not quite equal to that of the postrostral portion of the head.”

Gilbert and Hubbs (1920:388-390) recorded B. furvescens from Indonesia and the Philippines.
Howes and Crimmen (1990:195) erroneously referred these to B. cottoides (see Iwamoto and Merrett
1997:479 for comments on Howes and Crimmen’s treatment of B. furvescens). Our examination of
three of the five specimens (CAS-SU 25442, CAS-SU 25443, CAS-SU 25444) suggested that Gilbert
and Hubbs may have had a species different from B. furvescens. The three specimens were small and
in poor shape when examined, however, and certain of our measurements may not have been entirely
accurate. Notably, all three had 15 pyloric caeca, although Gilbert and Hubbs recorded 20 in one of

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 423

five specimens (they did not state which one). Additional material from the general region of the Phil-
ippines and Indonesia must be examined to properly determine which species actually occur there.

REFERENCE SPECIMENS. — AMS I.24355-016 (1 spec.); K83-18-02. AMS 1.25273-003 (1
spec.); K84-11-09. AMS 1.26001-016 (2 spec.); K86-01-09. AMS I.28070-002 (1 spec.); K88-11-02.
AMS 1.28372-002 (1 spec.); K88-08-05. AMS I.28712-002 (1 spec.); K88-10-02. AMS I.29338-002
(1 spec.); ORV Franklin stn FRO580-25, 28°05.76'S, 163°06.04'E; 1051 m; 5 May 1989. AMS
1.29812-002 (1 spec.); K89-15-02. AMS 1.30738-003 (1 spec.); K89-12-02.

REFERENCE. — Alcock (1894).

Bathygadus sp. cf. spongiceps Gilbert and Hubbs, 1920
Fig. 18

DISTINGUISHING FEATURES. — D II,8—10, P 114-118, V 9 (rarely 8 or 10); GR-I (outer)
(5—6)+19—20, total 25-26, GR-II (outer) 17—19; pyloric caeca 15—28, usually 20-28. Measurements
(in percent HL): snout length 30-33; internasal width 3 1-34; interorbital width 32—39; orbit diameter
16-22; suborbital width (bony) 12-16; postorbital length 52-55; distance orbit to preopercle 48-53;
length upper jaw 54—59; length pectoral fin 47—62; length pelvic fin 46-71; length outer gill raker
7—16. Outer pelvic ray in some specimens moderately prolonged, but most lack produced rays; pecto-
ral fin not extending beyond anus. Teeth bands relatively narrow to moderately broad, 7—12 teeth wide
at broadest part of premaxillary band, 4-8 wide in broadest part of dentary band. Fin rays dusky to
dark, but generally not black. Flesh and head bones rather soft and weak.

SIZE. — To approximately 50 cm.

DISTRIBUTION. — From most of the southwestern Pacific, New Zealand, Australia(NSW, WA),
Indonesia, and the Philippines. Depth range about 900-1500 m.

NSW CAPTURES. —A single specimen taken at each of the two deepest Kapala tows
(1130-1240 m), off Sydney and Jervis Bay; possibly more abundant at greater depths.

REMARKS. — Iwamoto and Williams (1999) found color differences in their material of B.
spongiceps from Western Australia, with some specimens considerably blacker than others, but they
found no other characters that would suggest specific differences. Merrett and Iwamoto (2000) exam-
ined specimens from the New Caledonian region that appeared to be identical to B. spongiceps except
for slight differences in the dentition of the lower jaw and counts of pyloric caeca. They called their
specimens B. sp. cf. spongiceps. We consider our NSW specimens as identical to the New Caledonian
species and have therefore followed Merrett and Iwamoto’s designation. More specimens of B.
spongiceps from the type locality must be examined and compared with specimens from other areas.

REFERENCE SPECIMENS. — AMS I.26001-005 (1 spec.); K86-01-09. AMS I.30394-03 (1 spec.);
K89-16-02.

REFERENCES. — Iwamoto and Merrett (1997)[in part; most specimens B. cottoides|; Iwamoto
and Williams (1999); Merrett and Iwamoto (2000).

Genus Gadomus

DISTINGUISHING FEATURES. — Gadomus and Bathygadus specimens are readily separated by
the former having a firmer body, stronger head bones and fin rays, and paler overall color of body. Al-
most all species of Gadomus have a long chin barbel, although two species have small or rudimentary
ones, and most have greatly elongated rays in one or more fins. Teeth are generally finer and in
broader bands in Gadomus.

REMARKS. — The taxonomy of the genus in the western Pacific and Indian Ocean is yet to be ad-
equately resolved. Howes and Crimmen (1990) distinguished two groups based on gill raker counts.

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Volume 52, No. 21

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

424

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 425

One of the New South Wales species falls in the group with low counts, the other (G. pepperi) in the
high-count group. Iwamoto and Williams (1999) provide a full account of both species.

Gadomus sp. cf. colletti Jordan and Gilbert, 1904
Fig. 19

DISTINGUISHING FEATURES. — D II,10; P 116-121; V 8; outer gill rakers short, about length of
gill filaments, (4—5)+(19-21), 24~25 total; pyloric caeca small, very numerous, more than 100. Barbel
thick, long, about 3 times diameter of orbit; gums and lower branchiostegal membrane pale; all fins
black; elongated dorsal ray less than twice head length, an extremely long upper pectoral ray, outer
pelvic ray about equal to or less than head length.

SIZE. — To 30cm.

DISTRIBUTION. — So far known only from Australia (NSW, WA), in 500-1150 m.

NSW CAPTURES. — Caught in a relatively narrow depth range of 690-975 m between Crowdy
Head and Batemans Bay, with most taken north of Sydney. Of the 65 specimens from 27 Kapala sta-
tions (15% of tows in 700—1000 m), 30 specimens were from two tows off Port Stephens; all other sta-
tions yielded fewer than five specimens per tow.

REMARKS. — Distinguished from G. pepperi by its lower gill raker count, shorter rakers on the
first gill arch, generally shorter elongated fin rays, and its pale mouth and tongue. Iwamoto and Wil-
liams (1999) discuss their reluctance to identify this species as G. colletti, which is described as hav-
ing a blackish buccal cavity, in contrast to the distinctly pale buccal cavity in the Australian
specimens.

REFERENCE SPECIMENS. — AMS 1.29813-006 (1 spec.); K89-06-05. AMS 1.24979-011 (1
spec.); K84-16-04. AMS I.19862-006 (1 spec.); K76-23-01. AMS 1.24659-001 (1 spec.); K84-06-04.
Others listed in Iwamoto and Williams (1999).

REFERENCE. — Iwamoto and Williams (1999).

FiGurE 19. Gadomus sp. cf. colletti Jordan and Gilbert, 1904. AMS 1.24659-001. From Kapala stn K84-06-04, off Broken
Bay, NSW, in 914-933 m.

426 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Gadomus pepperi lwamoto and Williams, 1999
Fig. 20

DISTINGUISHING FEATURES. — 1D II,9; P 114-118 (rarely 120); V 8; outer gill rakers long, twice
length of gill filaments, (5—6)+(22—25), total 28-31; pyloric caeca about 75 long, slender. Barbel slen-
der, length about one to two times orbit diameter; interorbital width 18-23% of HL; mouth all black;
gular membrane pale, branchiostegal membrane blackish, lips dark; fins dark but not intensely black,
tail tip dusky; first dorsal, pectoral, and pelvic fins with a prolonged ray generally exceeding twice
head length.

SIZE. — To 40 cm.

DISTRIBUTION. — Australia (Qld, NSW, west to WA), in 800-1500 m.

NSW _ CAPTUuRES. — Relatively common between Crowdy Head and Gabo Island in
825-1200 m. Recorded from 178 Kapala stations (including 94% of all tows deeper than 900 m); usu-
ally more than 10 specimens per tow.

REMARKS. — Distinguished from G. sp. cf. colletti by its higher gill raker counts, fewer pyloric
caeca, shorter barbel, dark mouth, pale gular membrane, longer fin rays, and less intensely black fins.

REFERENCE SPECIMENS. — AMS 1.24059-002 and 1.24059-020 (7 spec.); K83-09-02. Others
listed in Iwamoto and Williams (1999).

REFERENCE. — Iwamoto and Williams (1999).

Family Macrouridae

DISTINGUISHING FEATURES. — The most noticeable feature that separates this family from the
bathygadids is the protruding snout (the nasal bones have an anteriorly directed medial process that is
lacking in the bathygadids). Exceptions to this are Haplomacrourus nudirostris and Kuronezumia
spp., which may lack a protruding snout in adults, but unlike bathygadids they have small mouths. In
addition, almost all macrourids have spinules on the scales. The exceptions are some species of
Hymenocephalus, which may have lost the spinules secondarily.

REMARKS. — This large family contains about 300 species, many of which have yet to be de-
scribed. Three subfamilies are generally recognized, although each may warrant full family status.
Subfamily Macrourinae contains most of the 300 plus species in about 30 genera; Trachyrincinae has
about seven species in two genera; and Macrouroidinae has only two species in two genera.

REFERENCES. — Okamura (1970, 1989); Marshall (1973).

SUBFAMILY MACROURINAE

DISTINGUISHING FEATURES. — First dorsal fin high, the segmented rays preceded by one rudi-
mentary or spikelike spinous ray closely adpressed to a long spinous ray; first dorsal fin separated
from second dorsal by a distinct gap; anal fin rays much better developed (usually longer and stouter)
than those of second dorsal fin in almost all species. Branchiostegal rays 6—7 (rarely 8). Gill rakers
short, usually tubercular; the outer gill arch closely attached to gill cover by membrane, greatly reduc-
ing size of outermost gill slit.

427

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

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428 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

KEY TO GENERA AND SOME SPECIES OF MACROURINAE FROM NEW SOUTH WALES
(Adapted from Iwamoto and Williams 1999. Species in square brackets are not yet recorded from

NSW but can be expected.)

a. Second spinous ray, of first:dorsal fin‘smooti(Fig: 21a) ees 2 = re s- y Dy
1b. Second spinous ray of first dorsal fin serrated along leading edge (weakly or much reduced in some) (Fig. 21b). . . . 9
2a. Snout stoutly supported and pointed; a continuous suborbital ridge of coarsely spined scales extending from snout tip

to preopercle angle, terminating ina sharp point .......--.---++-++-+-++-+s-+: Caelorinchus (Fig. 22)
2b. Snout strongly to weakly supported, rounded to pointed; suborbital ridge, if present, not continuous from snout tip to

angle of preopercle and not terminating inasharppomt.........-.-+--+2++see+ + eae oe 3
3a. Broad areas of fine, parallel black lines (ventral striae) overlying silvery ground on ventral surfaces of chest, shoulder

sirdie, along each side ofasthmus, and’belly (Fig: 23; 24). 7... ee 4
3b: Noiventral'striae’. 22 BR cee k eas Se oe aeeeeere Sts Sle: hs eet. nee 5
4a. Ventral striae extend alongside anterior half or more of anal fin base; 6 branchiostegal rays; no lenslike light organ

onichestvattains more than O|cmiy lee) oe.) Seen Penne en Lepidorhynchus denticulatus (Fig. 23)
4b. Ventral striae rarely extend posterior to anus; 7 branchiostegal rays; a small lens of light organ on chest; adults small,

ustially less \thari.23) ermgh aati cthe sip sets ta: pa ee eRe 2 os seek Gop Dina et ela Hymenocephalus (Fig. 24)
5a. Lower jaw with large, widely spaced, fanglike teeth inonerow ....... 2... . 20 eee eee eee ee ees 6
Sb; Lower jaw with rather'small/teeth inmore'thanione TOW). =) as. ee a ee ee 8

6a. Head pores small; grooved lateral line complete to end of tail; chin barbel present; 7 branchiostegal rays
ee ES ee) ree eee 5 oo i oo O See neemurcmrenncrE Malacocephalus laevis (Fig. 25)
6b. Large open pores on head; grooved lateral line interrupted posterior to first dorsal fin; no chin barbel; 6 branchiostegal
TAYS ec a amie a eo wae aoa ho ke 5 GROMER coo oe =: Oe & Gicaen tn 7

7a. Anus removed from anal fin, usually about midway between pelvic fin and anal fin, preceded by a small black fossa
of light organ between pelvic fins; grooved lateral line single, short, terminating at vertical behind first dorsal fin;
abdomen short, distance isthmus to anus less than halfheadlength......... Odontomacrurus murrayi (Fig. 26)

FIGURE 21. Leading edge of second spinous ray of first dorsal fin smooth (a) and serrated (b).

suborbital ridge

FIGURE 22. Suborbital ridge in Caelorinchus extending from snout tip to preopercle angle, terminating in a sharp point.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 429

FiGuRE 23. Ventral striae in Lepidorhynchus denticulatus.

ventral striae

FIGURE 24. Ventral striae in Hymenocephalus sp.

FiGurE 25. Diagrammatic illustration of a Malacocephalus sp.

lateral line

FIGURE 26. Diagrammatic illustration of an Odontomacrurus murrayi showing position of anus and short lateral line.

430 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

7b. Anus immediately before anal fin, no fossa of light organ; grooved lateral line prominent, in two parts, an anterior
dorsolateral section and a posterior midlateral section; abdomen long, distance isthmus to anus more than

three-fourths ofiheadlengthiea: mecca ee tp mee ete i uy se ech srt Cynomacrurus piriei (Fig. 27)
Sar Pelvi¢TayS 0-7 reas cece ais chan Oe ee re een eis ee: aa ee Trachonurus (Fig. 28)
Sb: Pelvic trays 9512). sped Biansy eas Loe eee Get Oe cst ee fs eer [Ventrifossa sazonovi|
9a. Head massive, globose, soft; scales along base of second dorsal finenlarged...................... 10
9b. Head not especially massive or globose; no enlarged scales along base of second dorsal fin. ............. 1]

10a. Base of pelvic fin posterior to vertical through ongins of first dorsal and pectoral fins; interorbital width 31-34% HL
Pi ut ase GON Gc at AT eh naa net eeu RAE ee are CEE ah ah Sn hl [Cetonurichthys subinflatus]
10b. Base of pelvic fin about at or anterior to vertical through origin of first dorsal and pectoral fins; interorbital width

36=48 O/C (Rip 8 29) ae ee ce ee el ge eae Be ae Rn a oe a Cetonurus globiceps
lla. Scales of head elongated, with spinules longitudinally aligned to give striated pattern to head surfaces; chin barbel

ADSENE Mee eee Oy Bes eet ye tees pucks Me athe ly Cd AL A es ie Res Mesobius (Fig. 30)
11b. Head scales not elongated, no striated pattern to head surfaces; chin barbel present. ................. 12

12a. Snout rounded, not protruding beyond mouth in adults, naked; scales on head and front of body without spinules or
ridges; maxilla reaches only to vertical through front of orbit in adults (more posteriorly in juveniles); second spinous
ray of first dorsal fin notably large and laterally compressed, heavily serrated. . Haplomacrourus nudirostris (Fig. 31)
12b. Snout angular, protruding beyond mouth, completely naked to variously covered with scales; almost all scales
covered with spinules or low ridges; maxilla usually extends well posterior to front of orbit (except in some species
of Sphagemacrurus and Lucigadus); spinous second ray of first dorsal fin not greatly compressed laterally, sparsely

toidensely serrated/or smoothialonpileadmgiedge. = =~ | - erence | = eee 13
13a, Branchiostegal'tays'6) 25 6 ites Pe OR ee Oe a yw OM OA see cee 14
13bBranchiostegal.rayS'7) =. 5 Sc Foe ee ee A | aon Sew os Qe gay. ce cs eee 15
14a:Anusyatorcloseitojanalifin origin; nolightiorganl. =). 4+) -)2 4) 4) eee ee eee Coryphaenoides
14b. Anus removed from anal fin; small light organ present (Fig.32) .................. Mataeocephalus sp.

lateral line

FIGURE 27. Diagrammatic illustration of a Cynomacrurus piriei showing position of anus and lateral line in two parts.

enlarged scales

*iGURE 28. Diagrammatic illustration of a Trachonurus showing enlarged scales along base of anal fin.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 43]

FiGuRE 29. Diagrammatic illustration of a Cetonurus sp. (a) Lateral view showing relative positions of pelvic, pectoral, and
first dorsal fins. (b) Dorsal view of head; arrow points to enlarged scales along anterior part of second dorsal fin.

FiGuRE 31. Diagrammatic illustration of a Haplomacrourus nudirostris showing extensive naked areas on head (stippled)
and forward position of mouth.

branchiostegal rays

WS : a Mass SSOn
light organ anus

FIGURE 32. Diagrammatic ventrolateral view of Mataeocephalus sp. with six branchiostegal rays: anus far removed from
anal fin origin and small light organ before anus.

432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

15a. Snout mostly naked dorsally and ventrally without terminal and lateral tubercular scales... ............ 16
15b. Snout variously naked, but usually no extensive naked area dorsally; tubercular scales present at terminal and lateral
tipsiOl SnONE, Sas. geet sees ee) > cece ene et eee oP ee Sag inyea ceo) 18
16a. Olfactory organ huge, length of posterior nostril about one-half diameter of orbit . . . [Macrosmia phalacra] (Fig. 33)
16b. Olfactory organ normal, posterior nostril much less than half diameter of orbit... ................. 17
17a. Origin of anal fin below first dorsal fin; anus usually closer to pelvic fin than to anal fin....... [Kumba] (Fig. 34)

17b. Origin of anal fin well posterior to vertical through hind margin of first dorsal fin; anus immediately before anal fin
Dei Re ETE eh CCR eee DEMO EON RES oc ePIC Ee uae Asthenomacrurus victoris (Fig. 35)

18a. A double row of stout, modified scales under orbit forming a stout shelf and usually a sharp, rough ridge (Fig. 36) . 19

18b. Scales under orbit all small, forming smooth, rounded surface... 2... 2... 2... 2 eee ee ee ee 21
19a. Anus closer to pelvic fin insertions than to anal fin origin (Fig. 37a)... ............-2+2205, Nezumia
19b:Anus\closer to/anall fin than'toipelvic fin((kig. 3/D)=s— - 2 55 a cue = 4 ae ee ee 20

large posterior nostril

‘q{@~>
SS yy

FIGURE 33. Diagrammatic illustration of a Macrosmia phalacra showing extensive naked areas (stippled) on head and large
posterior nostril.

FiGURE 34. Diagrammatic illustration of a Kumba sp. showing extensive naked areas (stippled) on head and position of anal
fin origin relative to first dorsal fin.

FIGURE 35, Diagrammatic illustration of Asthenomacrurus victoris showing extensive naked areas (stippled) on head and po-
sition of anal fin origin relative to first dorsal fin.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 433

FIGURE 37. Diagrammatic ventral views of (a) Nezumia and (b) Sphagemacrurus showing positions of anus and relative
sizes of periproct region.

20a. Snout short, high, scarcely protruding beyond steeply oblique mouth; pelvic fin anteriorly placed, about under

preopercle; anal fin origin about under first dorsal finorigin......-.....--.--. Sphagemacrurus (Fig. 38)
20b. Snout long, protruding well beyond slightly oblique mouth; pelvic fin origin below or behind opercle, anal fin origin

below or behind vertical through posterior margin of first dorsal fin. ............ Mataeocephalus (Fig. 39)
21a. Outer gill rakers of second arch 13-18; no scales on gular and branchiostegal membranes . . . . Ventrifossa (Fig. 40)

21b. Outer gill rakers of second arch 12 or fewer; small scales sometimes present on gular or branchiostegal membranes 22

22a. Prominent fin markings (black blotches or streaks) in most species; spinules on body scales aligned in more or less
parallel rows; adult size less than 30 cm TL in most species... ...--.- 2-2-2 e es Lucigadus (Fig. 41)

22b. Fins lacking prominent markings; spinules on body scales in irregularly quincunx to somewhat divergent rows; adult
Cares erat tive | Ute ced tl Gene eet CRE enc! ae E SoNCmOn ae See Once mn armerececmr a Kuronezumia (Fig. 42)

434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

FiGurE 38. Diagrammatic illustration of a Sphagemacrurus sp. showing oblique mouth and relative positions of pelvic fin
and anal fin origin.

FIGURE 39. Diagrammatic illustration of a Mataeocephalus FIGURE 40. Diagrammatic illustration of a Ventrifossa sp.
sp. showing thick, coarse scales along suborbital and relative
positions of pelvic fin and anal fin origin.

FiGURE 41. Diagrammatic illustration of Lucigadus FIGURE 42. Diagrammatic illustration of Kuronezumia
microlepis. leonis.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 435

Genus Asthenomacrurus

DISTINGUISHING FEATURES. — Anus immediately in front of anal fin. Pelvic fin slightly in ad-
vance of vertical through pectoral fin base. Head bones weak, no coarse scaly ridges on head; snout al-
most entirely naked. Long spinous ray of first dorsal fin with few weak serrations along leading edge.
Seven branchiostegal rays. Light organ poorly developed or apparently absent. Species small, proba-
bly less than 25 cm.

REMARKS. — The genus is enigmatic and may eventually be considered the same as
Pseudonezumia Okamura, 1970. Only two species of small adult size, A. victoris, here reported, and
A. fragilis (Garman, 1899) from the eastern central Pacific. Specimens identified as Paracetonurus
sp. by Iwamoto (1986) are probably of this genus.

REFERENCE. — Sazonov and Shcherbachev (1982b).

Asthenomacrurus victoris Sazonov and Shcherbachev, 1982
Fig: 35

DISTINGUISHING FEATURES. — As for genus, with V7 or 8; outer GR-I 13-14.

SIZE. — Probably less than 25 cm.

DISTRIBUTION. — Indian Ocean, off Japan, and Australia(NSW, WA), in about 1650-3500 m.

NSW CAPTURES. — Two specimens were taken in 1650-1900 m off Nowra by the ORV Frank-
lin.

REMARKS. — The species was originally described from three specimens, one of which was
from about 200 n. mi. west of Freemantle (WA). The two NMV specimens are the shallowest captures
and the first record of the species from the Pacific coast of Australia.

REFERENCE SPECIMENS. — NMV A7000 (220 mm TL) and NMV A7001 (154 mm TL; 67 km
ene of Nowra (34°41.97'S, 152°22.44’'E); 1896-1642 m; ORV Franklin stn CSIRO FR9/88, Slope
59, 22 Oct. 1988.

REFERENCE. — Sazonov and Shcherbachev (1982b).

Genus Caelorinchus

DISTINGUISHING FEATURES. — Branchiostegal rays 6. A stout ridge formed of modified scales
extending from tip of snout to angle of preopercle, terminating posteriorly in a sharp point; other head
ridges stout in most species. Spinous ray of first dorsal fin smooth along leading edge. Pelvic fin rays
almost invariably 7. A ventral light organ with a black fossa either on belly or on chest in most species.
No gill rakers on outer side of first arch; rakers on inner side of arch tubercular, 10 or fewer total (ex-
cept in C. matamuus, with as many as 12). Chin barbel present.

REMARKS. — This is the most speciose genus of grenadiers with more than 100 known and more
awaiting description. The genus is best represented in tropical and subtropical waters, but is also nu-
merous in certain temperate waters such as off New Zealand, where 21! species have been recorded.
We have found 16 species of Caelorinchus off New South Wales. Most of these are widespread in the
southern part of the continent, with other species more representative of the subtropical-tropical
Queensland fauna.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto (1990); McMillan and Paulin (1993);
Iwamoto and Williams (1999).

436 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

KEY TO THE SPECIES OF CAELORINCHUS OF NEW SOUTH WALES

la. Underside of snout naked --:--:+:-:::*++-:-:: D
1b. Underside of snout fully scaled- -----------: 8
2a. A prominent black fossa of light organ on midline of
chest or belly (Fig. 43) oh Ny eno Sone nne Romane c 3
2b4Nonossaionchestonbellys 6a a ene 7

3a. Rays of second dorsal fin long, about equal to opposite
rays of anal fin; black fossa of light organ on chest - -
2 Qe 299 DDD Geng cia of C. sp. cf. cingulatus (Fig. 44)
3b. Rays of second dorsal fin short, much shorter than oppo-
site rays of anal fin; black fossa of light organ on belly4

light organ

4a. A small but prominent black spot at base of pectoral fin;
pelvic fin with large black blotch in middle of fin; light
organ large, extending anteriorly to or beyond trans-
verse line connecting origins of pelvic fins (Fig. 45): -
Se iol te, Nap Wien efecey Jn) Goi ciel te) teeter sec a! Metale tot tnteereMol ce Mar vey “a (G. mirus

4b. No spot at base of pectoral fin; pelvic fins lacking large
black blotch; light organ extends forward no further
than to line connecting insertions of pelvic fins: - - - 5

anus

FiGuRE 43. Ventral view of trunk of a Caelorinchus show-
5a. Dorsally behind leading edge of snout with usually clear _ing large anterior dermal window of light organ between pel-
naked area on each side of midline (Fig. 46); saddles vic fins.
usually absent on trunk, but faintly present on tail; pale
interspaces between saddles spotlike in dorsal view
(Fig. 47)
Sa), URE Wea VAS ate oS UPSAET. 5) SU eae MO CL peda Sov Fie; cet ead Salar gi eager een at 9 aoe C. parvifasciatus
5b. Dorsally behind leading edge of snout densely covered with scales, lacking clear naked areas; saddles prominent on
trunk/and'tail® nterspaces not spotlike inidorsaliview) = =) 30) 5) es 6

6a. First dorsal and anal fins black or very dark to base, without prominent pale areas (Fig. 48); body scales relatively
deciduous; pyloric'caecal2—=23;\(Fig3 49). 0.6 ec ee he Pe ey Se ee C. fasciatus

6b. Base of first dorsal fin distinctly pale, anal fin with a prominent black stripe posteriorly (Fig. 50); body scales relatively
adherentjpyloricicaccayl8—34 (usually 22-32)" 3) ec) a cee ee C. maurofasciatus

FIGURE 44. Diagrammatic illustration of Caelorinchus sp. cf. cingulatus showing pigmentation pattern and high second dor-
sal fin.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

437

nara Et

light organ

anus

pan eens ome ==

FIGURE 45. Ventral view of a Caelorinchus mirus showing

FiGurE 46. Dorsal view of snout of Caelorinchus
large anterior dermal window of light organ and prominent
black spot on pelvic fins.

parvifasciatus showing naked areas behind leading edge.

pale, spotlike

FIGURE 47.
marks on tail.

pyloric caeca

FIGURE 48. Diagrammatic illustration of Caelorinchus

FIGURE 49. Ventrolateral view of a Caelorinchus showing
fasciatus showing banding pattern and pigmentation of first
dorsal and anal fins.

cut-away of left abdominal wall exposing stomach and pyloric
caeca.

FiGureE 50. Diagrammatic illustration of Caelorinchus maurofasciatus showing banding pattern and pigmentation on first
dorsal and anal fins.

438 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

7a. Trunk completely encircled with broad bluish band (Fig. 51); pyloric caeca 10—14; anterolateral margin of snout
incompletely;supported by, bone (E1272) anwar nen enn ee ee C. kaiyomaru

7b. Trunk not encircled by bluish band; pyloric caeca 7-9; anterolateral margin of snout completely supported by bone
Teer eres 8.6 es Se er ce i eee ent Mec ga o 6 c C. innotabilis

8a. Body with about 8-10 pale longitudinal stripes; light organ relatively large, extending forward to about midpoint
between pelvic and anabfinibases. 4. 2 5 eg sa ss gS ches ao S oer eer C. australis
8b. Body lacking longitudinal stripes; light organ small, usually not extending to midpoint between anal and pelvic fins . 9

9a. Anterior one-half to one-third of anal fin black, remainder pale; trunk completely encircled by dark band; orbit equal

to\or longemthan snopt length, cs cba ae ec vel nine) st coe deen se C. matamuus (Fig. 53)
9b. No sharp contrast in anal fin pigmentation; trunk not encircled by dark band; orbit diameter much shorter than snout

length, 3s Ste ey eel ens Sete Whee ce eso nd sie of ole ae Rh 10
10a. Anterolateral snout margin completely supported by bone; spinule rows on body scales 3-8, widely divergent

(Pig; 54a):) ogee hS 0a se eo we See lle ins Ress San, RR 11
10b. Anterolateral snout margin incompletely supported by bone; spinule rows on body scales 3—13, more or less parallel

(Fig. S4b) se we foe ke ed) eee Bees A aiea, Fe, Egy dO ce) 42s en 13

medial process

gap

lateral process

FiGuRE 51. Diagrammatic illustration of Caelorinchus
kaiyomaru showing broad dark band encircling trunk.

FIGURE 52. Diagrammatic ventral view of head of a
Caelorinchus sp. showing partial cut-away of ventral snout
surface to expose lateral and medial processes of nasal bone (a
wide gap between processes).

FIGURE 53. Diagrammatic illustration of Caelorinchus FiGuRE 54. Diagrammatic illustrations of Caelorinchus
matamuus showing broad dark band encircling trunk and _ body scales taken from region below anterior end of second
black anterior portion of anal fin. dorsal fin. (a) C. smithi—spinule rows widely divergent, (b)

C. mycterismus—spinule rows more or less parallel.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 439

11a. Snout length in adults 1.5—1.7 times orbit diameter (1.7—2.0 in young); GR-II (total inner) 6—7; scale rows below
Sige MMMATSUIOLSAI TEN O= Oi aed 5-5 hte) ee takoes, os Hest 09. Sn eee AER easy & Sat see aia ns ge eae C. smithi
11b. Snout length 1.7—2.5 times orbit diameter; GR-II (total inner) 8—10; scale rows below origin of first dorsal fin 6-8. 12

12a. GR-I 8 total; scales rows below origin of first dorsal fin 6—7, lateral line scales over a distance equal to predorsal

length 38-44; orbit diameter 1.27—1.44 into postorbital length... ..........-...-...- C. macrorhynchus
12b. GR-I 9-10 total; scales rows below origin of first dorsal fin 8, lateral line scales over a distance equal to predorsal
length 53-55; orbit diameter 0.97—1.08 into postorbital length .................--.. C. supernasutus
13a. Scale rows below origin of first dorsal fin 8.0—10.0, below second dorsal fin 5.0—7.5; nasal fossa usually naked,
RCP SCAlea in Sone (Pigs S5a)e shift Neale PLP. LPP ee eds PES) IETS oe AY. C. acanthiger
13b. Scale rows below origin of first dorsal fin 4.0-7.0, below second dorsal fin 3.5—7.0; nasal fossa naked or scaled
Mem OD) ss 8 2 es able wis, 5) Ae Spe Re ay ws RS ede 2s oy CPD AT se Gest ee 14
14a. Scales on head ridges notably coarse and sharply spined; 5—13 parallel rows of spinules on body scales (Fig. 56a);
overall color dark grayish with violet tinge; fins all blackish. ... 2... 2.2. ee eee ee ees C. trachycarus
14b. Scales on head not especially coarse; 4-9 more or less parallel rows of spinules on body scales (Fig. 56b); overall
color tawny to grayish brown, not tinged with violet; fins dusky to blackish... . 1... 2-2-2 225-554. 15

15a. Snout 1.9-2.2 into HL; anal fin dusky except pale near posterior tip; snout slightly upturned (Fig. 57a) C. mycterismus
15b. Snout 2.42.6 into HL; anal fin dusky to blackish overall; snout lacking upturned tip (Fig. 57b). . . . C. kermadecus

FIGURE 56. Scales from region below anterior end of second
dorsal fin of (a) Caelorinchus trachycarus and (b) C.
kermadecus.

FIGURE 57. Lateral view of snout of (a) Caelorinchus
mycterismus showing upturned tip and (b) C. kermadecus
with straight tip.

440 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Caelorinchus acanthiger Barnard, 1925
Fig. 58

DISTINGUISHING FEATURES. — 1D II,8—9; Pil7-119; V 7; GR-I 8—9 total; scales below 1D 8-10,
below 2D 5-7.5, lat.l. about 33-49; pyloric caeca 9-12. Snout about 2/5ths of head length, with
straight to slightly concave dorsal profile; anterolateral margin incompletely supported by bone. Un-
derside of head covered with small scales; nasal fossa usually entirely naked; body scales with 3-5
parallel spinules rows, middle row longest, other rows usually much shorter. Small black fossa imme-
diately anterior to anus.

SIZE. — To 50cm.

DISTRIBUTION. — Southern Africa, southern Australia (NSW, Vic., Tas., SA, WA), and New
Zealand in mid-slope depths of about 800 m to at least 1200 m.

NSW _ CAPTuRES. — Relatively abundant between Crowdy Head and Gabo Island in
790-1200 m. Caelorinchus acanthiger was recorded from 224 Kapala stations, including 66% of
tows in 800—900 m and all tows deeper than 900 m. Between 10 and 50 specimens were captured in
most trawls.

REMARKS. — Among grenadiers, C. acanthiger is second only to Corvphaenoides serrulatus in
abundance in mid-slope depths off NSW. The species is distinguished from two similar species, C.
kermadecus and C. mycterismus, by its higher scale-row counts below the first dorsal fin and by the
absence of scales on the nasal fossa (present ventrally in other two species). C. mycterismus also has a
noticeably longer snout. C. trachycarus has much more spiny scales, especially on head ridges, more
spinule rows on body scales, a darker overall color, black fins, and fewer scale rows below the first
dorsal fin. C. macrorhynchus, C. smithi, and C. supernasutus have the anterolateral snout margins
completely supported by bone.

REFERENCE SPECIMENS. — AMS I.19860-017 (1 spec.); K76-24-03. AMS 1.20068-010 (5
spec.); K77-23-13. AMS I.20098-023 (1 spec.); K77-23-07. AMS 1.20099-018 (3 spec.); K77-23-12.
AMS 1.21722-002 (1 spec.); K79-20-13. AMS I.21724-011 (1 spec.); K79-20-15. AMS 1.24037-012
(6 spec.); K78-26-16. AMS 1I.24054-004 (1 spec.); K83-06-02. AMS 1.24056-007 (2 spec.);
K83-08-02. AMS 1I.24059-007 (1 spec.); K83-09-02. AMS 1I.24060-007 (1 spec.); K83-09-01. AMS
1.24100-001 (6 spec.); K83-07-11. AMS 1.24157-006 (1 spec.); K83-12-04. AMS 1.25273-007 (1
spec.); K84-11-09. AMS I.28475-002 (1 spec.); K88-04-06. AMS I.28745-004 (1 spec.); K88-04-06.
AMS 1.28749-002 (2 spec.) and AMS I.28749-003 (1 spec.); K88-17-03. AMS I.29798-01 1 (4 spec.);
K89-12-04. AMS I.29807-002 (1 spec.); K89-07-05.

REFERENCES. — Trunov (as C. pseudoparallelus) (1983); Gomon et al. (1994); Iwamoto and
Anderson (1994); Iwamoto and Williams (1999).

Caelorinchus australis (Richardson, 1839)
Fig. 59

DISTINGUISHING FEATURES. — ID II, 9-11; P il3-118; V 7; GR-I 7—9 total; scales below 1D
3.5—5.5, below 2D 3.54.5, lat.1. about 24-32; pyloric caeca 31—34. Snout about one-third of head
length; anterolateral margin incompletely supported by bone. Underside of head covered with small
scales; nasal fossa with small scales over ventral surfaces; body scales with as many as 20 parallel
rows of short, small spinules. Large black fossa of light organ extends forward from anus about half-
way to pelvic fin insertions. About 8—10 pale longitudinal stripes on dorsolateral aspects of body; first
dorsal fin black on distal half; anal fin dusky to blackish along distal margin, mostly blackish posteri-
orly. (Adapted from Arai and McMillan 1982)

SIZE. — To at least 55 cm.

44]

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

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442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

DISTRIBUTION. — Southeastern Australia (NSW, Vic., Tas., SA) on shelf and upper slope in
about 100-450 m. Off Tasmania, Last et al. (1983) stated that the species “is commonly trawled be-
tween 80 and 300 metres but on rare occasions has been collected in shallow coastal waters.”

NSW CAPTURES. — Caught by Kapala only south of Batemans Bay in depths of 130-440 m.
Moderately abundant, C. australis was caught in 22% of all tows in 100-400 m but was most common
in 200-300 m (present in 55% of tows in that depth range). This species of relatively large adult size
has no commercial value, but its numbers off southern NSW appear to have been reduced by trawling
over the last 20 years. During 1976-77 it was recorded in 80% of Kapala tows in 200-400 m, and
catches were mostly greater than 20 specimens per tow; in comparison, it was present in only 23% of
1996-97 tows, with an average of 3 per tow.

REMARKS. — The species name was previously applied to a common grenadier of New Zealand
waters until Arai and McMillan (1982) determined that C. australis was confined to southeastern
Australia. The New Zealand species for which the name was being applied 1s endemic and was undes-
cribed until Arai and McMillan named it. Note that fig. 21.12 in Last et al. (1983) is of C.
biclinozonalis.

REFERENCE SPECIMENS. — AMS I.29385-002 (2 spec.); K89-03-15.

REFERENCES. — Arai and McMillan (1982); Last et al. (1983); Gomon et al. (1994)

Caelorinchus sp. cf. cingulatus Gilbert and Hubbs, 1920
Fig. 60

DISTINGUISHING FEATURES. — 1|D II, 8—9; P 116-119; V 7; GR-I 6-8 total; scales below 1 D 5-7,
below 2D 4.5-—6.0, lat.1. about 30-40. Snout two-fifths to one-half of head length; anterolateral margin
incompletely supported by bone. Underside of head naked; nasal fossa naked or sparsely scaled; body
scales with short, spikelike, recumbent spinules in 7—15 parallel to slightly divergent rows. Light or-
gan extends forward from anus to chest, dilated at each end, anteriorly with a shallow, scaled fossa.
Rays of second dorsal fin about as high as opposites of anal fin; interspace between first and second
dorsal fins short, usually less than length base of first dorsal. A saddle extending from base of first dor-
sal to base of pectoral fin; a second faint saddle usually visible below origin of second dorsal fin; a
darker third saddle below 9th—12th rays of second dorsal extending anteroventrally and leveling off
midlaterally; faint saddles posteriorly on body, but usually not extending ventrally below lateral line;
first dorsal with a dark midlateral band.

SIZE. — To about 30 cm.

DISTRIBUTION. — New Caledonia, and ne. Australia (Qld, NSW). Depth range about
250-550 m.

NSW CAPTURES. — Five specimens collected by Kapala from three stations in 550 m off the
QlId-NSW border (about 28°S).

REMARKS. — Iwamoto and Merrett (1997) first reported this species as C. cingulatus from speci-
mens collected in the New Caledonian region. In their subsequent study (Merrett and Iwamoto 2000),
they realized that their specimens, though closely similar, did not entirely agree with descriptions of
C. cingulatus Gilbert and Hubbs, 1920, especially in regards to certain body markings, but they did
not describe it as new. The species is readily distinguished from other NSW members of the genus by
the combination of its distinctive pattern of saddles; moderately prolonged spinous first dorsal ray,
high second dorsal, the rays about equal in length to opposites of anal fin; and relatively long span be-
tween the isthmus and anal fin origin.

REFERENCE SPECIMENS. — AMS ]1.20518-012 (2 spec.); K78-09-05. AMS I.20459-014 (3 spec.)
and 1.20459-019 (1 spec.); K78-17-10. AMS I.20651-014 (1 spec.); K78-23-09.

REFERENCES. — Iwamoto and Merrett (1997); Merrett and Iwamoto (2000).

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 443

Caelorinchus fasciatus (Giinther, 1878)
Fig. 61

DISTINGUISHING FEATURES. — ID II,9—10; P 114-118; V 7; GR-I 8—9 total; scales below 1D
3.5—5.0, below 2D 3.5-4.5, lat.l. about 21-29; pyloric caeca 12—23. Snout 18-33% of head length,
much shorter than huge orbit, which is about two-fifths or more of head length; anterolateral margin of
snout incompletely supported by bone. Underside of head naked; dorsal surface of snout lacking clear
scaleless areas; nasal fossa with small scales over ventral surfaces. Body scales large, rather decidu-
ous, exposed fields covered with 7—12 parallel rows of spinules. Narrow fossa of light organ extends
forward from anus about halfway to pelvic-fin insertions. A series of 8—12 dark saddle marks on body
beginning on nape. First dorsal fin black on distal half or more; anal fin dusky to blackish along distal
margin, mostly blackish posteriorly.

SIZE. — To at least 35 cm.

DISTRIBUTION. — Southeastern Australia (NSW, e. Vic.), New Zealand, and South America.
Depth range off NSW about 600—1000 m, much more restricted than the 200—1000 m off New Zea-
land (P. McMillan, pers. commun. with Graham). Off Chile and Argentina, Iwamoto (unpublished re-
cords) has verified records only from much shallower depths of about 75-450 m.

NSW CAPTURES. — Uncommon off NSW; 19 specimens captured in eight tows by Kapala in
630-960 m, from just south of Sydney (34°15’S) to Gabo Island (37°40’S).

REMARKS. — It is likely that the Kapala specimens treated here are the only Australian records
definitely referable to C. fasciatus. Australian records of C. fasciatus in the literature prior to about
1990 most probably relate to C. maurofasciatus or C. parvifasciatus (e.g., Last et al. 1983; Munro
1957). In addition to NSW, Vic. and Tas., Last et al. (1983: 240) reported C. fasciatus from WA and
SA, but Iwamoto and Williams (1999) found no specimens to verify those records. The species was
not treated in The fishes of Australia’s south coast (Gomon et al. 1994). Records of C. fasciatus off
southern Africa were based on different species (see Iwamoto and Anderson 1994).

Caelorinchus fasciatus and C. maurofasciatus are closely similar and difficult to differentiate.
Characters that appear to separate the two species include the presence in C. fasciatus of one or two
large, thick, somewhat elevated scales on the median line of the nape two to four scales forward of the
first dorsal fin. The pyloric caeca count also differs: 12-23 (x = 16.6) in C. fasciatus (data from
McMillan and Paulin 1993), compared with 18-34 in C. maurofasciatus (McMillan and Paulin
[1993] gave 22-32, x = 27.6). Compared to C. maurofasciatus, the overall body color of C. fasciatus,
including banding, is duller, the anal fin lacks a dark stripe, the first dorsal fin is uniformly dusky to
blackish, and the scales are more deciduous.

REFERENCE SPECIMENS. — AMS 1.24774-001 (3 spec.); K84-08-05. AMS 1.26998-003 (1
spec.); K87-14-02. AMS 1I.32431-002 (1 spec.); K89-07-04.

REFERENCES. — Last et al. (1983); McMillan and Paulin (1993).

Caelorinchus innotabilis McCulloch, 1907
Fig. 62

DISTINGUISHING FEATURES. — 1D IT,9—10; P 116-119 V 7; GR-I 6-8 total; scales below 1D 6-7,
below 2D 6.0-—7.5, lat.l. about 32-40; pyloric caeca 7—9. Snout slender and sharp, two-fifths or more
of HL, much longer than orbit diameter, which is about one-third of HL; anterolateral margin sharp,
completely supported by bone. Underside of head naked anteriorly, but some small, nonimbricate
scales in small patches above and behind mouth; nasal fossa naked; body scales with 9—13 parallel
rows of short, slender spinules. Light organ externally inconspicuous, not generally visible without
dissection, relatively short, extending forward from anus to level of pelvic fin insertions; anus re-

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IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 445

moved from anal fin origin by 2 or 3 scale rows. No distinct markings on body or fins; second dorsal
fin rays about as long anteriorly as anal fin rays.

SIZE. — To about 35 cm.

DISTRIBUTION. — Southern Australia (NSW, Vic., Tas., SA. WA) and New Zealand.

NSW CAPTURES. — Captured by Kapala between Crowdy Head and Gabo Island in
450-1075 m. Recorded from 227 tows (about half of all tows between 500 and 1100 m), with greatest
abundance in 700—900 m (in 93% of tows at that depth range). Frequently more than 50/tow in trawls
with small-meshed codends.

REMARKS. — This is a plain fish with no distinguishing pigment pattern, an almost-cylindrical
body, and a sharp suborbital ridge that is completely supported by bone along the leading edge of the
snout. These features separate C. innotabilis from most other NSW members of the genus (C. smithi
and C. macrorhynchus also have complete support of the leading snout margin). Caelorinchus
innotabilis might be confused with C. kaiyomaru, but the dark, broad band girdling the trunk in C.
kaiyomaru is distinguishing.

REFERENCE SPECIMENS. — AMS 1.7893 (holotype, 138 mm TL); off Sydney, 800 fm [1463 m;
but see note in Historical Perspective]; Woy Woy. AMS 1.15973-010 (1 spec.); K71-07-03. AMS
1.15976-001 (1 spec.); K71-09-01. AMS 1.16589-003 (1 spec.); K72-05-05. AMS 1.18726-027 (5
spec.); K75-01-02. AMS I.18770-009 (2 spec.); K75-02-08. AMS I.18838-010 (1 spec.); K75-05-03.
AMS I.18839-001 (28 spec.) and AMS I.18839-037 (25 spec.); K75-05-04. AMS 1.19198-002 (2
spec.); K76-05-04. AMS I.19202-002 (1 spec.); K76-06-03. AMS 1.19859-002 (10 spec.);
K76-24-04. AMS I.19860-009 (1 spec.); K76-24-03. AMS I.19862-008 (2 spec.); K76-23-01. AMS
1.20098-011 (2 spec.); K77-23-07. AMS 1I.20452-015 (1 spec.); K75-05-05. AMS 1.21722-008 (3
spec.); K79-20-13. AMS I.21724-013 (1 spec.); K79-20-15. AMS I.21806-001 (3 spec.); K77-07-10.
AMS 1.23885-014 (1 spec.); K78-27-05. AMS 1.24055-006 (2 spec.); K83-08-01. AMS I.24056-005
(6 spec.); K83-08-02. AMS 1.24059-008 (4 spec.); K83-09-02. AMS 1.24060-005 (1 spec.),
1.24060-008 (3 spec.), and 1.24060-016 (2 spec.); K83-09-01. AMS 1.29756-005 (1 spec.);
K89-15-04. QM 1.23010 (1 spec.); e. of Terrigal, NSW, in 446 fm; Oct. 1978.

REFERENCES. — Gomon et al. (1994); Last et al. (1983); Iwamoto and Williams (1999).

Caelorinchus kaiyomaru Arai and lwamoto, 1979
Fig. 63

DISTINGUISHING FEATURES. — | DII,7—10; Pi16—i20, V 7; GR-I 6-9 total; scales below 1D 5-8,
below 2D 5.0-6.5, lat.l. about 31-41, usually 35-40; pyloric caeca 10—14. Snout slender and sharp,
two-fifths to almost half of head length, much longer than orbit diameter (25-30% of HL);
anterolateral margin incompletely supported by bone. Underside of head naked or with | or 2 isolated
scales above posterior end of mouth; nasal fossa naked; body scales with 8-10 parallel or slightly di-
vergent rows of small spinules. Light organ short, seen only as a small blackish area before anus; anus
removed from anal fin origin by | or 2 scale rows. Entire trunk encircled by broad dark, bluish band;
blackish orbital ring.

SIZE. — To 40 cm.

DISTRIBUTION. — Australia (NSW, Vic., Tas.), New Zealand, South Atlantic off Falkland Is., in
about 845—1150 m.

NSW CAPTURES. — Caught by Kapala in 880-1150 m on all grounds south of Crowdy Head.
Relatively common in mid-slope depths and recorded from 148 stations (present in 76% of all tows
between 900 and 1100 m), frequently more than 10 specimens per tow.

REMARKS. — This slender species is readily recognized by the prominent dark blue color com-
pletely encircling the trunk. In NSW Caelorinchus species, this character is found only in C.
matamuus, a large-sized, heavy-bodied species.

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IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 447

REFERENCE SPECIMENS. — AMS 1.24055-002 (1 spec.); K83-08-01. AMS 1.24059-005 (7
spec.); K83-09-02. AMS I.24060-012 (3 spec.), I.24060-015 (2 spec.); K83-09-01. AMS I.24150-007
(1 spec.); K83-13-01. AMS 1.24157-005 (1 spec.); K83-12-04. AMS 1.24173-008 (3 spec.);
K83-14-06. AMS 1I.24355-009 (2 spec.); K83-18-02. AMS 1.24356-001 (1 spec.); K83-14-05. AMS
1.24462-002 (1 spec.); K83-15-02. AMS I.24565-001 (1 spec.); K83-14-03. AMS 1.24980-005 (4
spec.); K84-16-15. AMS 1.29737-005 (1 spec.); K89-19-01.

REFERENCES. — Arai and Iwamoto (1979); Gomon et al. (1994).

Caelorinchus kermadecus Jordan and Gilbert, 1904
Fig. 64

DISTINGUISHING FEATURES. — 1D II,7—9; P 116-119; V 7; GR-I 7-9 total; scales below 1D
4.5—6.0, below 2D 4.5-6.0, lat.1. about 32-38; pyloric caeca 10—12. Snout 2.4—2.6 into head length in
adults (longer in smaller specimens); anterolateral margin incompletely supported by bone. Under-
side of head covered with small scales; nasal fossa finely scaled ventrally; body scales with 4—7
more-or-less parallel rows of broadly triangular spinules, middle row largest. Light organ short, not
externally visible. Overall color grayish brown; all fins dusky to blackish; mouth dark gray to
blackish; area around anus bluish, but color not extending to bases of pelvic fins.

SIZE. — To about 60 cm.

DISTRIBUTION. — Kermadec Is., New Zealand, s. of New Caledonia, and Australia (NSW), in
about 800—1150 m.

NSW CAPTURES. — Only seven specimens taken at five Kapala stations between Crowdy Head
and Batemans Bay.

REMARKS. — The Kapala specimens are the only Australian records of this species, but it can be
expected off Queensland. Caelorinchus kermadecus is most similar to C. acanthiger, differing princi-
pally in squamation features. Body scales are larger (fewer scale rows below the dorsal fins) than in C.
acanthiger, and the nasal fossa is scaled ventrally (naked in C. acanthiger).

REFERENCE SPECIMENS. — AMS 1.24991-003 (1 spec.); K84-16-05. AMS 1.28100-001 (1
spec.); K88-08-06. AMS I.29750-001 (1 spec.), AMS I.29750-004 (1 spec.), and AMS 1.29750-005
(1 spec.); K89-17-04. AMS 1.29798-013 (1 spec.); K89-12-04. AMS 1I.29807-003 (1 spec.);
K89-07-05.

REFERENCES. — McMillan and Paulin (1993); Iwamoto and Merrett (1997).

Caelorinchus macrorhynchus Smith and Radcliffe, 1912
Fig. 65

DISTINGUISHING FEATURES. — 1D II,8—9; Pil6—19; V 7; GR-I 8-10 total; scales below 1 D 6-7,
below 2D 5.5, lat.l. about 38-44. Snout length 47-50% HL; interorbital width 19-20%; suborbital
width 11-12% HL; postorbital length (30-31%); length orbit to angle of preopercle (32-34%); upper
jaw length (24-25%), outer gill slit length 1 1-13%; and barbel length (8—9%). Snout narrow, sharply
pointed, about one-half of head length; anterolateral margin completely supported by bone. Under-
side of head covered with small scales; nasal fossa mostly covered with small scales; body scales with
5-8 divergent rows of broad-based spinules. Light organ small, a black streak extends forward from
ventral fossa to midway between anus and pelvic fin base. Overall color swarthy to black; mouth and
fins blackish.

SIZE. — To about 50 cm.

DISTRIBUTION. — Philippines, Indonesia, and Australia (Qld, NSW, WA), in about
500-1100 m.

NSW CAPTURES. — Uncommon; taken only six times (7 specimens) by Kapala between New-
castle (33°S) and Jervis Bay (35°S), in 550-950 m. One AMS specimen (1.26806-003) was collected

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IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 449

by acommercial trawler in 550 m off Port Stephens. A juvenile (CAS 214042, 145+ mm TL) was col-
lected recently off Montague I. (36°12’S) in 500 m.

REMARKS. — A notably long snout, dark color, and divergent spinule rows distinguish
Caelorinchus macrorhynchus from most other NSW members of the genus. The species resembles C.
supernasutus in its notably long snout, small orbits, and small ventral light organ. Caelorinchus
supernasutus, however, has smaller scales (8 rows below 1D origin, 5.5 below mid-base of 1D, and
53-55 lateral line scales over a distance equal to predorsal length), and shorter measurements of sub-
orbital (9—10% HL), postorbital (24-25%), orbit to angle of preopercle (26%), upper jaw (20-21%),
and barbel (46%). One specimen (AMS I.29601-001) listed under this species has a noticeably short
(1.5 times orbit) broad snout, very unlike that in most other specimens examined, and more like that in
C. smithi. However, scale features distinguish that specimen from C. smithi, and other characters ap-
pear to be the same as those of C. macrorhynchus. Because of the short snout in the specimen, it will
not key out properly in the key to species. More specimens are needed to adequately delimit the range
of variation in snout length and shape in C. macrorhynchus.

REFERENCE SPECIMENS. — AMS I.21722-007 (1 spec.); K79-20-13. AMS 1.24625-005 (1
spec.); K84-06-06. AMS 1I.24778-001 (2 spec.); K84-08-02. AMS 1.26806-003 (1 spec.); FV
Vincenzann; e. of Port Stephens, 550 m; 25 Oct. 1986. AMS I.29600-001 (1 spec.); K87-24-01. AMS
1.29601-001 (1 spec.); K88-08-08. AMS I.29825-001 (1 spec.); K89-09-09. CAS 214042 (1 spec.);
FV Shelley H, off Montague I. (36°12'S, 150°24’E); 490-525 m; 1 Mar. 2000.

REFERENCES. — Radcliffe (1912); Iwamoto and Williams (1999).

Caelorinchus matamuus (McCann and McKnight, 1980)
Fig. 66

DISTINGUISHING FEATURES. — 1D II,8—10; P 116-119; V 7; GR-I 12-13 total; scales below 1D
7-10, below 2D 7-9; pyloric caeca 18-29. Snout bluntly conical, anterolateral margin not completely
supported by bone; mouth large, upper jaw extends to below middle of orbit; orbit about one-third of
HL. Underside of head scaled; head ridges stout but not especially spiny; nasal fossa covered with
scattered small scales; body scales with 8-11 slightly divergent rows of low spinules. Light organ
small, immediately before anus, not externally visible. Trunk completely encircled by a broad,
blue-black band; anterior half to one-third of anal fin black, remainder pale.

SIZE. — To about 65 cm.

DISTRIBUTION. — Southeastern Atlantic to southern Africa, across Indian Ocean to southern
coast of Australia (NSW, Vic., Tas., SA, WA), and New Zealand, in about 650-1100 m.

NSW CAPTURES. — Caught frequently in small numbers by Kapala in depths between 690 and
1010 m onall grounds south of Broken Bay (33°25’S). Overall, about 200 specimens captured in 44
Kapala trawls; most frequently caught in 700-900 m, where it was recorded in 44% of all tows in that
depth range.

REMARKS. — Caelorinchus matamuus is a distinctive, widespread species of the Southern
Ocean having a geographical distribution similar to that of C. acanthiger. The large size, broad bulky
head, blunt snout, prominent blackish trunk band, and black anterior part of anal fin immediately iden-
tify the species.

REFERENCE SPECIMENS. — AMS I.18726-024 (2 spec.); K75-01-02. AMS 1.19860-012 (1
spec.); K76-24-03. AMS I.20099-005 (1 spec.), 1.20099-009 (1 spec.), I.20099-017 (1 spec.);
K77-23-12. AMS I.20485-015 (1 spec.); K77-23-06. AMS I.23885-005 (1 spec.); K78-27-05. AMS
1.24054-002 (3 spec.); K83-06-02. AMS I. 24157-007 (1 spec.); K83-12-04. AMS 1.24613-006 (1
spec.); K75-05-05.

REFERENCES. — McCann and McKnight (as Mahia matamua) (1980); Sazonov and Shcher-
bachev (1982a); Last et al. (1983); Iwamoto and Williams (1999).

450 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Caelorinchus maurofasciatus McMillan and Paulin, 1993
Fig. 67

DISTINGUISHING FEATURES. — 1D II,9-10; P il5—i19; V 7; GR-I 7-9 total; scales below 1D
4.56.0, below 2D 4-5, lat.I. about 22-31; pyloric caeca 18-34. Snout about one-third or less of head
length, much less than huge orbit; anterolateral snout margin incompletely supported by bone. Under-
side of head naked; nasal fossa partially scaled; no broad naked areas above leading edge of snout;
body scales with 11 or more rows of short, small spinules. Slender fossa of light organ between pelvic
and anal fins. About 9—1 1 prominent saddle marks, the first beginning forward on nape and ending be-
low anterior portion of first dorsal fin, the second saddle beginning under posterior end of first dorsal
fin and ending two scale rows behind origin of second dorsal fin; most fins dark distally; first dorsal fin
blackish distally, with paler base; anal fin with dark stripe; mouth dark.

SIZE. — To at least 50 cm.

DISTRIBUTION. — Southern Australia (NSW, Vic., Tas., SA, WA) and New Zealand, in about
300-900 m.

NSW CAPTURES. — Absent off northern NSW, but recorded south of about 32°20’S in
320-820 m. Very abundant on upper slope, particularly in 400-700 m. During 1996-97, C.
maurofasciatus was caught in 74 Kapala tows in 400-650 m (94% of all tows in that depth range) be-
tween Sydney and Gabo Island, with an average catch of 57/tow.

REMARKS. — Caelorinchus maurofasciatus is most similar to C. fasciatus, with which it has
been confused (see description of that species for comparison). The prominent black saddle markings
and the black stripe along the anal fin distinguish the species from similar NSW members of the genus.

REFERENCE SPECIMENS. — AMS I.15970-005 (1 spec.); K71-06-04. AMS 1I.18839-014 (10
spec.), AMS 1.18839-016 (8 spec.); K75-05-04. AMS 1.19197-001 (1 spec.); K76-04-03. AMS
1.23470-006 (5 spec.); K82-17-01. AMS 1.23862-001 (10 spec.); K81-18-05. AMS 1.24854-003 (1
spec.); K84-14-01. AMS I.28713-001 (1 spec.); K84-13-03. NMV A2460 (1 spec.); K81-17-03.

REFERENCES. — McMillan and Paulin (1993); McMillan in Gomon etal. (as Caelorinchus sp. 1,
the “false banded whiptail”) (1994); Iwamoto and Williams (1999).

Caelorinchus mirus McCulloch, 1926
Fig. 68

DISTINGUISHING FEATURES. — 1D IT,9—10; P il7—i19; V 7; GR-I 7-8 total; scales below 1D
5.0-5.5, below 2D 4.0-5.5, lat.l. about 23-26; pyloric caeca about 40. Snout less than one-third of
head length, much less than large orbit; anterolateral snout margin incompletely supported by bone.
Underside of snout and suborbital anteriorly naked, lower jaw and preopercle scaled; nasal fossa and
lunate areas above leading edge of snout naked; body scales with 9-20 rows of small spinules. Naked
fossa of light organ large, extends to origin of pelvic fin. No saddle markings except in juveniles
(<100 mm TL) with banding pattern; pectoral fin with small black spot at ventral corner, pelvic fin
with black blotch; other fins dusky to pale, but anal fin with blackish distal margin posteriorly.

SIZE. — To about 30 cm.

DISTRIBUTION. — Australia (Qld, NSW, e. Vic., WA) in about 100-500 m.

NSW CAPTURES. — One of the most abundant species of genus and found along entire coast of
NSW in depths between | 10 and 500 m. Recorded from 293 Kapala stations, with most tows at depths
between 200 and 400 m. Despite the large codend mesh (90 mm) used during 1996-97 surveys, C.
mirus was caught in 43 tows between 200 and 350 m (65% ofall tows in that depth range), with amean
catch of 153/hour tow; some individual catches were in excess of 1000 fish. Caelorinchus mirus is one
of only two NSW grenadiers (also C. australis) inhabiting outer shelf depths as well as the slope. It is

451

IWAMOTO AnD GRAHAM: GRENADIERS OF NEW SOUTH WALES

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FiGuRE 68. Caelorinchus mirus McCulloch, 1926. AMS 1.26221-002. From Kapala stn K85-20-10, off Port Stephens,
NSW, in 154-157 m.

caught at night by prawn trawlers off northern NSW in depths as shallow as 110 m(K. Graham, pers.
observ.).

REMARKS. — Caelorinchus mirus is endemic to Australia. It has been erroneously recorded off
New Zealand, but McMillan and Paulin (1993) have determined that it does not exist there. There are
no confirmed records of C. mirus from Tas., western Vic., or SA. This apparent disjunct distribution
between the east coast and WA (including the Great Australian Bight) is similar to those of Gadomus
sp. cf. colletti (see above) and a number of endemic Australian sharks and rays (see Last and Stevens
1994).

REFERENCE SPECIMENS. — AMS I.19205-004 (4 spec.); K76-07-01. AMS 1.21793-010 (2
spec.); K78-17-11. AMS 1.23993-008 (2 spec.); K78-17-14. AMS I.25932-006 (1 spec.); K85-21-06.
AMS 1.26221-002 (3 spec.); K85-20-10. Others listed in Iwamoto and Williams (1999).

REFERENCES. — McMillan and Paulin (1993); McMillan in Gomon et al. (1994); Iwamoto and
Williams (1999).

Caelorinchus mycterismus McMillan and Paulin, 1993
Fig. 69

DISTINGUISHING FEATURES. — 1D II,7—9; P 116-119; V 7; GR-I 6-8 total; scales below 1D 4—5,
below 2D 5-7; pyloric caeca 8-11. Snout 1.9-2.2 into head length in adults (longer in smaller speci-
mens), upturned anteriorly; anterolateral margin incompletely supported by bone. Underside of head
covered with small scales; nasal fossa finely scaled; body scales with 49 parallel to slightly divergent
rows of triangular spinules, middle row highest and longest, lateral rows lower and shorter. Light or-
gan short; no black, lens-like fossa. Overall color in alcohol tawny to brownish; all fins dusky except
posterior part of anal fin pale; mouth dark; area around anus bluish.

SIZE. — To about 50 cm.

DISTRIBUTION. — New Zealand, s. of New Caledonia (on Norfolk Ridge), and Australia (NSW,
possibly WA) in about 850-1150 m.

NSW CAPTURES. — Two juveniles, one taken east of Crowdy Head in 1050 m, the other off
Nowra in 950-978 m.

REMARKS. — New South Wales is probably outside its normal range, but the species may be ex-
pected off Qld. Two juveniles of less than 25 cm TL were tentatively recorded by Iwamoto and Wil-
liams (1999) from the North-West Shelf off WA. C. mycterismus bears close resemblance to several

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 453

FIGURE 69. Caelorinchus mycterismus McMillan and Paulin, 1993. AMS 1.24057-004. From Kapala stn K83-09-04, east of
Nowra, NSW, in 951-978 m.

other members of the genus from the southwestern Pacific, including C. acanthiger, C. trachycarus,
and C. kermadecus. The first two are immediately distinguished by their naked nasal fossae (among
other characters). The longer snout with slightly upturned tip and the pale posterior end of the anal fin
generally suffice to distinguish C. mycterismus from C. kermadecus.

REFERENCE SPECIMENS. — AMS 1.24057-004 (1 spec.); K83-09-04. AMS 1.29750-007 (1
spec.); K89-17-04.

REFERENCES. — McMillan and Paulin (1993); Iwamoto and Williams (1999).

Caelorinchus parvifasciatus McMillan and Paulin, 1993
Fig. 70

DISTINGUISHING FEATURES. — 1D II,9—12 (usually 10); P il7—i20; V. 7; GR-I 7-8 total; scales
1D 5-6, 2D 4.5-5.5, lat.I. 26-28; pyloric caeca 19-26. Snout short, broad, blunt; anterolateral margins
incompletely supported by bone; orbit longer than snout length. Naked fossa of light organ extends
forward close to line connecting pelvic fin insertions. Underside of head naked or with I—3 small
scales above angle of lower jaw; broad, naked, translucent areas dorsally behind leading edges of
snout; nasal fossa naked. Seven or eight faint, sometimes almost obscure, saddles posteriorly on body,
more pronounced posteriorly; narrow pale bands occupying one or two scale rows at dorsomedial line
separating saddle marks, pale areas often with appearance of white dorsal spots; mouth and gill cavi-
ties dark.

DISTRIBUTION. — Southeastern Australia (NSW, Vic., Tas.) and New Zealand.

NSW CAPTURES. — Commonly captured by Kapala between Crowdy Head and eastern Bass
Strait in depths of 220-600 m; a single record off the Clarence River. This species of relatively small
size is possibly the most abundant Caelorinchus off central and southern NSW and was recorded from
more than 250 Kapala upper-slope stations. During the 1996-97 Kapala survey, 95 trawls (75% of to-
tal) in 250-550 m averaged almost 200 C. parvifasciatus per tow, despite the 90 mm codend mesh.

SIZE. — To about 30 cm.

454 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 70. Caelorinchus parvifasciatus McMillan and Paulin, 1993. AMS 1.26240-001. From Kapala stn K85-17-02, east
of Broken Bay, NSW, in 421—457 m.

REMARKS. — Our Australian specimens agreed in most respects with the New Zealand
type-specimens (see McMillan and Paulin, 1993, from which we gleaned the following comparative
data). We noted some differences in the ranges of certain measurements and in the count of pectoral
fin rays (11 7—i20 in Australia cf. 115-118 in types); snout length 27-31% of HL cf. 29-34%; preoral
length 26-34% cf. 21-35%; orbit diameter 44-49% cf. 31-46%; suborbital width 16-18% cf.
13-16%; upper jaw length 26-33% cf. 20-30%; and barbel length 9-14% cf. S—11%. The anterior
dermal window of the light organ was generally larger in New Zealand specimens, but there was over-
lap in the proportional measurements of that length. Our specimens were also darker than described
for the species, especially the fins. The first dorsal fin is dark, almost blackish with a pale base, com-
pared with dusky in the type-specimens; the pectoral fins are dark dusky to blackish, compared with
pale; the pelvic fins are blackish with the outer ray distally white, compared with “dusky blackish”;
and the anal is blackish overall, paler posteriorly, compared with “pale with a dusting of
melanophores diffuse anteriorly.” Finally, our specimens completely lacked the one to three small
scales on the ventral surface of the head above the articulation of the lower jaws, and the count of py-
loric caeca in 31 of our specimens ranged somewhat lower than that given in the original description
(19-26, x = 21.8, compared with 22—28, x = 25.3).

Caelorinchus parvifasciatus closely resembles C. mirus, but its relatively small light organ dis-
tinguishes it from that species. It differs from C. fasciatus and C. maurofasciatus in having less promi-
nent saddle marks, spotlike pale markings along the dorsal midline of the tail, and broad naked areas
behind the leading edge of the snout. The last two species also attain a much larger size than does C.
parvifasciatus.

A NSW specimen of C. parvifasciatus (AMS 1.15975-036) was misidentified by Iwamoto and
Williams (1999:128) as C. amydrozosterus. That species has a different banding pattern without
spotlike pale interspaces, a slightly larger dermal window of the light organ, and fewer pyloric caeca.
Caelorinchus amydrozosterus has yet to be recorded from the Australian east coast, although it is
commonly caught with C. parvifasciatus off Portland, western Victoria (K. Graham, pers. observ. ).

REFERENCE SPECIMENS. — AMS 1I.15968-013 (5 spec.); K71-05-04. AMS I.15970-027 (4
spec.); K71-06-04. AMS 1.15973-008 (3 spec.); K71-07-03. AMS 1.15994-007 (35 spec.);
K71-13-02. AMS I.16565-003 (1 spec.); K72-04-01. AMS I.18774-001 (1 spec.); K75-03-02. AMS
1.18838-017 (19 spec.); K75-05-03. AMS I.18839-054 (1 spec.); K75-05-04. AMS I.19205-003 (4
spec.); K76-07-01. AMS 1.20301 -006 (1 spec.); K77-13-12. AMS I.24127-001 (1 spec.); K75-05-02.
AMS 1.26240-001 (1 spec.); K85-17-02.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 455

REFERENCES. — McMillan and Paulin (1993); McMillan in Gomon et al. (as Caelorinchus sp. 3,
“little whiptail’’) (1994).

Caelorinchus smithi Gilbert and Hubbs, 1920
Fig: 71

DISTINGUISHING FEATURES. — 1D II,8—10; 115-119; V 7; GR-I 7-8 total; scales below 1D
5.0-6.0, below 2D 4.5—6.0, lat.1. about 29-37; pyloric caeca 19-26. Snout 1.9—2.4 into head length;
anterolateral margin completely supported by bone. Underside of head covered with small scales; na-
sal fossa finely scaled anteriorly and ventrally to almost naked; body scales with 3—7 divergent rows
of stout, triangular spinules, middle row strongest, with 4—6 spinules; all rows complete to edge of
scale. Light organ short, externally visible as short black fossa before anus. Overall color dark brown
to swarthy; all fins blackish; mouth blackish; belly region bluish.

SIZE. — To about 35 cm.

DISTRIBUTION. — Philippines to Indonesia and Australia (Qld, NSW, NT), in about 400—750 m.

NSW CAPTURES. — Only one specimen caught by Kapala in 740 m off Qld-NSW border.

REMARKS. — Caelorinchus smithi is a tropical species and may be confused in NSW waters only
with C. macrorhynchus, which has a longer snout (about 2 in HL cf. 2.0—2.3 in C. smithi), smaller orbit
(4-5 in HL cf. 3.54.2), and somewhat more gill rakers on first arch (8—10 cf. 7-8).

REFERENCE SPECIMEN: AMS I[.21795-012 (1 spec.); K78-23-08.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto and Williams (1999).

FIGURE 71. Caelorinchus smithi Gilbert and Hubbs, 1920 (from Iwamoto and Williams 1999:fig. 22).

456 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Caelorinchus supernasutus McMillan and Paulin, 1993
Fig. 72

DISTINGUISHING FEATURES. — 1D II,9; P 118-19; V 7; GR-I 9-10 total; scales below 1D 8, be-
low 2D 6.5, lat.l. about 53-55. Snout length 52-54% HL; interorbital width 17—18%; suborbital width
10-11% HL; postorbital length (24-25%); length orbit to angle of preopercle (26%); upper jaw length
(20-21%), outer gill slit length 9%; and barbel length (4-6%). Snout long, narrow, tipped with a
broad, flat diamond-shaped scute; length about one-half of head length; anterolateral margin com-
pletely supported by bone. Underside of head covered with small scales; nasal fossa mostly covered
with small scales; body scales with 5—8 divergent rows of broad-based spinules. Light organ small but
externally visible with black dermal window before anus. Overall color darker dorsally and blackish
over abdomen and gill covers; lips, gums, mouth lining dark; ridge of median nasal bone dark; first
dorsal fin blackish, other fins pale to dusky.

SIZE. — To about 64 cm.

DISTRIBUTION. — Australia (NSW) and New Zealand, in about 500—900 m.

NSW CAPTURES. — Taken twice by Kapala, off Crowdy Head and Nowra, in 500-900 m.

REMARKS. — The small size and few captures suggest that NSW is outside the normal range of
this primarily New Zealand species. In having a long snout, small orbit, and small dermal window of
the light organ immediately before the anus, Caelorinchus supernasutus most closely resembles C.
macrorhynchus (the two species are compared in the description of C. macrorhynchus). Our two
NSW specimens of C. supernasutus are juveniles of 162 and 280 mm TL; they were not compared
with the much larger (418-635 mm TL) paratypes from New Zealand. Specimens of comparable size
from the two areas should be compared to verify our identification.

REFERENCE SPECIMENS. — AMS I.27609-001 (1 spec.); K87-23-02. AMS 1.29738-001 (1
spec.); K87-24-05.

REFERENCE. — McMillan and Paulin (1993).

Caelorinchus trachycarus lwamoto, McMillan, and Shcherbachev, 1999
Fige73

DISTINGUISHING FEATURES. — 1D II,7—9; P i15-i18; V 7; GR-I 7-9 total; scales below 1D
4.5-7.0, below 2D 3.5-6.0, lat.1. about 28-37; pyloric caeca 7-9. Snout about 2.0—2.5 of head length,
with straight to slightly concave dorsal profile; anterolateral margin incompletely supported by bone.
Underside of head scaled; nasal fossa naked to suborbital ridge; head ridges especially spiny and
coarse; body scales with 5—13 parallel spinules rows, middle row longest and highest, other rows usu-
ally much shorter and lower. Light organ small, not externally visible.

SIZE. — To about 50 cm.

DISTRIBUTION. — Southern Australia (NSW, Vic., Tas., SA, WA), New Zealand, and Norfolk
Ridge s. of New Caledonia, in 622—1730 m.

NSW CAPTURE: Only one confirmed specimen from off Jervis Bay in 1130 m.

REMARKS. — Caelorinchus trachycarus is relatively more abundant in deeper, more southern
waters of New Zealand and Australia, especially in the Great Australian Bight. Almost all reported
captures were from depths greater than 1000 m. It is likely to be confused with the common C.
acanthiger, which has smaller scales, more spinules rows on body scales, weaker spines on head
ridges, paler body and fin color, and absence of a violet tinge to the body.

REFERENCE SPECIMEN: AMS I.28475-004 (1 spec.); K88-04-06.

REFERENCES. — Iwamoto et al. (1999); Iwamoto and Williams (1999).

457

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

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458 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Genus Cetonurus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Head massive, globose, with capacious,
fluid-filled chambers; head ridges lacking thick modified scales. First dorsal fin short, high, with steep
base; slightly prolonged spinous ray serrated along leading edge. Anus surrounded by broad, circular
naked area, the anterior edge of which extends most of distance between anal and pelvic fins. Scales
small, densely covered with short, erect spinules; head entirely scaled, including branchiostegal mem-
branes and usually gular membrane. Grooved lateral line interrupted, not continuous; scales along,
and anterior to, second dorsal fin base enlarged, usually with enlarged spinules.

REMARKS. — Two widespread species: C. globiceps, the only species found in NSW, and C.
crassiceps (Giinther, 1878).

REFERENCE. — Sazonov and Shcherbachev (1985).

Cetonurus globiceps (Vaillant in Filhol, 1884)
Fig. 74

DISTINGUISHING FEATURES. — ID II,7—11, usually 9-10; P 115-119; V 8—11, usually 9-10;
GR-I 10-14; pyloric caeca 7—11. Orbit diameter 24-32% of HL; interorbital width 36-48%.

SIZE. — To about 40 cm.

DISTRIBUTION. — Widespread in central Atlantic, southern Africa, Indian Ocean, Australia
(NSW, Vic., SA, WA), New Zealand, and Japan.

NSW _ CAPTURES. — Caught by Kapala between Crowdy Head and Batemans Bay in
940-1200 m. A total of 99 specimens was caught in 34 tows (19% of all tows deeper than 900 m).

REMARKS. — Sazonov and Shcherbachev (1985) provided important information on the two
species of the genus. Cetonurus globiceps can be distinguished from C. crassiceps by the former hav-
ing a larger orbit, narrower interorbital, and somewhat more scale rows below first dorsal fin (13-19
vs. 11-14).

REFERENCE SPECIMENS. — AMS 1.24057-002 (1 spec.); K83-09-04. AMS 1.24187-001 (1
spec.); K83-14-02. AMS 1.24355-006 (3 spec.) and AMS I.24355-011 (3 spec.); K83-18-02. AMS
1.24624-002 (1 spec.); K84-04-10. AMS 1.25273-002 (6 spec.); K84-11-09. AMS 1.29605-005 (2
spec.); K89-09-07.

REFERENCES. — Sazonov and Shcherbachev (1985); Gomon et al. (1994); Paxton et al. (1989) as
C. crassiceps.

FIGURE 74. Cetonurus globiceps (Vaillant in Filhol, 1884). AMS 1.24057-005. From Kapala stn K83-09-04, east of Nowra,
NSW, in 951-978 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 459

Genus Coryphaenoides

DISTINGUISHING FEATURES. — Branchiostegal rays 6. Anus at or close to anal fin origin; no as-
sociated light organ. Spinous second ray of first dorsal fin serrated along leading edge (sometimes ru-
dimentary or lost). Rays of second dorsal fin much shorter than opposite rays of anal fin. Outer gill
rakers present (sometimes rudimentary) on first gill arch.

REMARKS. — The members of this genus are mostly found at mid-slope to lower-slope depths,
but a few range down to abyssal levels. The genus is represented in all ocean basins, from polar to
equatorial seas. Of the more than 60 species found worldwide, 11 were captured off NSW, with
Coryphaenoides serrulatus one of the most abundant of all grenadiers. In contrast, the deepest-living
Coryphaenoides species were represented by only one or a few specimens, probably reflecting the
paucity of sampling in depths greater than 1200 m.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto (1990); Iwamoto and Shcherbachev
(1991). Shcherbachev and Iwamoto (1995).

KEY TO THE SPECIES OF CORYPHAENOIDES FROM NSW

la. Pelvic fin rays 7, rarely 6 or 8; spinules on body scales lanceolate... ............-.....--220-- 2
1b. Pelvic fin rays 8 or more; spinules on body scales needlelike................-..--.---5-++-055- 3

2a. Chin barbel rudimentary; a greatly elongated ray in pectoral fin; inner gill rakers on first arch 16-19 . C. subserrulatus
2b. Chin barbel well developed, 20-30% of head length; no elongated ray in pectoral fin; inner gill rakers on first arch

ISLES IES MPMI tre ro fe ec ee he sates eae. sich a) PES MPa set inp sic ssirsapascnen eee rate, hare Ene C. serrulatus
3a. Snout completely scaled or naked surfaces confined to midventral swath and along ventral snout margin ...... . 4
3b. Snout entirely or almost entirely naked ventrally (and often on dorsal surface) ..............-..-...--. 7
4a. Outer gill slit greatly restricted, 4-9% of head length; inner rakers on first gill arch9 or 10 total ........ C. rudis
4b. Outer gill slit 14% of head length or greater; inner rakers on first gill arch 11 or moretotal............... 5
awe ViCHMirAaySIOwEALCLY HOLS paste c aiuse) 1 pape Sis > Ged coos as ews oks &@ © Ghee a er Gel eos C. dossenus
Oh, Ses pCa DN oe ee ne te ee re eee ne ee ec es 6

6a. Preopercle with 4 spikelike struts (Fig. 75); head about 6 times into total length; teeth rather small and weak,
SOMeWNalGeClIGUOUSNP Etsy fist tal eee, fo te or Skee Oe es Meee alc tect Mea OO Ce cos asa Mec aes C. grahami
6b. Preopercle lacking spikelike struts; head about 5.5 times into total length; teeth strong, tightly attached . . C. striaturus

7a. Chin barbel less than 5% of head length, usually a mere stump; pelvic fin rays 8, rarely 9; inner gill rakers on outer arch
IG=H1O) oe J bia ie A ah Se eR EE ee Ao ce Mono aceon C. memillani

8a. Inner gill rakers on outer arch 14—16; chin barbel 23-26%
of head length; outer gill slit 20-22% of head length -
5 di (nS ate Beh el ee ee C. murrayi
8b. Inner gill rakers on outer arch 9-13; chin barbel 4-16%
of head length; outer gill slit 12-19% of head length: 9

9a. Long spinous ray of first dorsal fin with numerous
serrations along leading edge; outer gill rakers on first
arch 6— 1 3 Sibchien io oes uw uacteo@Nee ew sh auch emeliiemidr s,s! ce) -suhe! as 10

9b. Long spinous ray of first dorsal fin with few or no
serrations along leading edge; outer gill rakers on first
ae Sls Goo Whe lo. G6) coro bel aac C. filicauda

10a. Snout distinctly pointed in lateral view, protruding be-
yond mouth a distance about equal to half orbit diame-
ter; chin barbel 9-16% of head length - - C. carapinus

10b. Snout blunt, scarcely protruding beyond mouth, ventral

profile steep; chin barbel short, 5—9% of head length
Peri. Pe Rom SAA C. sp. cf. fernandezianus FiGURE 75. Preopercle of Coryphaenoides grahami
showing spikelike struts.

spikelike struts

460 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Coryphaenoides carapinus (Goode and Bean, 1883)
Fig. 76

DISTINGUISHING FEATURES. — ID IT,8—9; P 116-120; V 9-10; GR-I (total, outer/inner series)
6-9/11—13; pyloric caeca 6-9. Snout length 33-37% of HL; orbit diameter 17—21%; interorbital
width 34-39%; suborbital width 1 2—16%; upper jaw length 35-40%. Head about 5—6 in TL; chin bar-
bel short, thin, 9-13% of HL. Dentition in upper jaw three or four teeth across at widest portion, outer
series slightly enlarged; teeth in lower jaw in one row except at symphysis in about two rows. Outer
gill rakers of first arch short, flaplike. Head lacking stout ridges of modified scales; leading edge of
snout with row of small, deciduous, tubercular scales; snout otherwise naked. Body scales large, de-
ciduous; spinules on exposed field reduced or lacking. Long spinous ray of first dorsal fin with numer-
ous short, reclined spinules along leading edge. Pectoral fin relatively long, about 50-70% HL. Outer
pelvic ray slightly produced, its length 50-80% HL. Color in alcohol overall gray to brownish, some-
what darker on head; blackish over operculum, gill membranes, and jaws; fins pale except for blackish
spinous ray of first dorsal, uppermost ray of pectoral, and outer ray of pelvic.

SIZE. — To about 40 cm.

DISTRIBUTION. — Worldwide at bathyal depths; recorded around southern Australia (NSW,
Vic., Tas., SA, WA) in 1000-3000 m, but range to 4900 m in eastern North Atlantic.

NSW CAPTURES. — A single NSW record (6 specimens) by ORV Franklin off Nowra in
1600-1900 m.

REMARKS. — Probably occurs too deep to have been sampled by Kapala. The description above
for Coryphaenoides carapinus is based on Australian specimens only. Specimens from other areas
differ from Australian specimens in certain counts and measurements, and we are uncertain if these
differences are indicative of separate taxa or populations.

REFERENCE SPECIMENS. — NMV A7003 (6 spec., 85-254 mm TL); 67 km off Nowra, NSW,
34°41.97'S, 151°22.44'E, in 1896-1642 m; ORV Franklin stn CSIRO FR9/88, Slope 59; 22 Oct
1988.

REFERENCES. — Marshall and Iwamoto (1973); Haedrich and Polloni (1976).

Coryphaenoides dossenus McMillan, 1999
Fig. 77

DISTINGUISHING FEATURES. — 1D IT,9—10; P 117-121; V 8 (rarely 7 or 9); GR-I (total, outer/in-
ner series) 7—9/1 1-13, GR—II 9-12/11-—13; scales below 1D 9.5—11, below 2D 7-12, lat.1. 38-48; py-
loric caeca 10-18. Snout length 25-29% HL; orbit diameter 19-23%; interorbital width 16-20%;
suborbital width 9-13; upper jaw length 40-45%. Head long, shallow, its width about half its length,

If
Wi SS
Onn Se
Sve ’ »\ = Se a
epee
ayn / \
r f/ \

FIGURE 76. Coryphaenoides carapinus (Goode and Bean, 1883). CAS 58671, from eastern Indian Ocean, about 300 n.mi. w.
of Perth, Western Australia.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 461

FIGURE 77. Coryphaenoides dossenus McMillan, 1999. (a) Female, AMS 1.24658-001, from Kapala stn K84-08-03, east of
Nowra, NSW, in 869-924 m; (b) male, AMS 1.26245-015, from Kapala stn K86-01-07, off Sydney, NSW, in 819-899 m.

about 4.2—5.6 in TL (in females, more in males); snout low, blunt, barely protruding; chin barbel well
developed, its length 21-34% HL. Upper jaw teeth in broad band, with outer series enlarged and
widely spaced; lower jaw teeth in 3 or 4 irregular rows laterally. Gill rakers somewhat tablike. Head
ridges not especially prominent and not reinforced by enlarged, thickened scales; underside of snout
mostly covered with small scales, although narrow ventral margin naked; body scales rather large,
with numerous subparallel to slightly convergent rows of small, needlelike spinules. Tip of outer pel-
vic ray barely or not reaching anal fin origin. Color in alcohol variable from light brown to swarthy
overall, fins dusky to blackish, mouth and gill cavities dark.

SIZE. — Males to about 50 cm; females to more than 85 cm.

DISTRIBUTION. — Widespread around New Zealand, New Caledonia and the Coral Sea, Austra-
lia (Qld, NSW, Vic., Tas., SA, WA) and in the Indian Ocean; also in the southeastern Atlantic from
South A frica north to the Gulf of Guinea. Depth range from about 700 to 1600 m, but most commonly
around 900—1200 m.

NSW CAPTURES. — Captured frequently but in small numbers on all mid-slope grounds be-
tween Crowdy Head and Gabo Island in depths from 695 to 1200 m. It was present in 143 Kapala
trawls including 51% of those deeper than 700 m; average catch about 3 per tow.

REMARKS. — Females (Fig. 77a) are more robust and attain a much larger size than males (Fig.
77b); larger individuals generally have a well-marked humped nape. Specimens of C. dossenus are

462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 2]

unlikely to be mistaken for other members of the genus owing to the combination of low, scarcely pro-
truding snout, large mouth, large size, few pelvic fin rays, relatively high number of gill rakers, and
long barbel.

REFERENCE SPECIMENS. — AMS 1.17866-007 (1 female); K72-07-01. AMS I.17867-007 (2 fe-
males); K72-07-04. AMS 1I.18726-026 (1 female); K75-01-02. AMS 1.19859-012 (1 female);
K76-24-04. AMS I. 20477-002 (1 female); K77-23-10. AMS 1.20485-006 (1 female), AMS
1.20485-011 (1 female); K77-23-06. AMS 1I.21724-003 (7 males), AMS 1.21724-006 (1 male);
K79-20-15. AMS I. 24037-006 (3 females); K78-26-16. AMS 1.24055-012 (1 female); K83-08-01.
AMS 1.24056-004 (2 females); K83-08-02. AMS 1.24059-016 (2 females); K83-09-02. AMS
1.24173-015 (1 female); K82-14-06. AMS 1.24613-005 (1 female), AMS 1.24613-007 (3 females);
K75-05-05. AMS 1.24624-001 (3 males), AMS 1.24624-006 (2 females); K84-04-10. AMS
1.24658-002 (1 female); K84-08-03. AMS I.24771-002 (1 female); K84-10-08. AMS 1.24980-006 (1
female); K84-16-15. AMS 1.24981-003 (2 females); K84-17-04. AMS 1.24992-002 (2 females);
K84-11-07. AMS I.25273-005 (1 female); K84-11-09. AMS 1.25415-001 (1 female); K84-04-11.
AMS 1.25933-005 (1 female); K79-20-06. AMS I.26245-003 (2 females) and 1.265245-015 (1 male);
K86-01-07. AMS I.28717-003 (1 female); K88-10-04. NMV A6842 (2 spec.); 56 km off Nowra,
NSW, 34°44.0’S, 151°14.5'E, in 817-1009 m; ORV Franklin stn CSIRO FR9/88, Slope 58; 22 Oct
1988.

REFERENCES. — McMillan in Gomon et al. (1994); Shcherbachev and Iwamoto (1995);
McMillan (1999).

Coryphaenoides sp. cf. fernandezianus (Ginther, 1887)
Fig. 78

DISTINGUISHING FEATURES. — 1D II,8; P il7; V 9-10; GR-I (total, outer/inner series)
10-11/11—12. Snout length 27-36% of HL; preoral length 16-18% of HL; orbit diameter 18—20%;
interorbital width 31-37%; suborbital width 15—16%; upper jaw length 40-41%. Head about 5 in TL;
chin barbel short, thin, 5—9% of HL. Dentition in upper jaw in two series; teeth in lower jaw in one
row. Outer gill rakers of first arch short, flaplike. Head lacking stout ridges of modified scales; few
small, deciduous scales on suborbital and lower jaw, snout naked ventrally and along anterior part of
dorsal surface. Body scales deciduous; spinules on exposed field reduced or lacking. Long spinous ray
of first dorsal fin with numerous short, reclined spinules along leading edge. Outer pelvic ray slightly
produced, its length about 70% HL. Color in alcohol overall dark brownish, somewhat darker on
head; blackish over operculum, gill membranes, and jaws; fins dark.

SIZE. — To at least 16 cm.

DISTRIBUTION. — Australia (NSW and Lord Howe Rise) in 1600-2500 m.

NSW CAPTURES. — Two juveniles captured by ORV Franklin, one off Nowra in about 1800 m,
the other on the Lord Howe Rise in 2450 m.

REMARKS. — We are uncertain of the identity of these two small specimens, but many of their
characters are similar to those of C. fernandezianus (Giinther, 1887), a species known only from the
holotype taken off Juan Fernandez Island off the west coast of Chile. Our two specimens differ suffi-
ciently from one another that they may represent separate species. The Nowra specimen has a longer
snout (36% of HL vs. 27% in the Lord Howe Rise specimen), broader interorbital (37% of HL vs.
31%), shorter orbit to preopercle distance (48% HL vs. 54%), and fewer pelvic fin rays (9 vs. 10).
More specimens are needed to resolve the identification questions.

REFERENCE SPECIMENS. — AMS _1.29316-005 (1 spec.); Lord Howe Rise, 29°42.06’S,
159°48.31'E; 2450 m; ORV Franklin, 3 May 1989. NMV A7002 (1 spec.); 67 km off Nowra,
34°41.97'S, 151°22.44’E, 1896-1642 m; ORV Franklin, Slope 59, stn FR 9/88, 22 Oct. 1988.

REFERENCES. — Giinther (1887); Iwamoto and Sazonov (1988).

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 463

FiGuRE 78. Coryphaenoides sp. cf. fernandezianus. AMS 1.29316-001. From ORV Franklin stn off Lord Howe Island, in
2450 m.

Coryphaenoides filicauda Ginther, 1878
Fig. 79

DISTINGUISHING FEATURES. — 1D II,8—10; P 116-119; V 9-10; GR-I (total, outer/inner series)
3-6/9-10; pyloric caeca 6-8. Snout length 33-38% of HL; orbit diameter 16—18%; interorbital width
30-33%; suborbital width 12-17%; upper jaw length 33-37%. Head about 5—6 in TL; chin barbel
short, thin, 4-12% of HL. Upper jaw teeth in narrow band, with slightly enlarged outer series; lower
jaw teeth in narrow band tapering to one row posteriorly. Outer gill rakers of first arch few, weakly de-

FIGURE 79. Coryphaenoides filicauda Giinther, 1878. AMS 1.27643-004. From Kapala stn K88-12-02, southeast of Crowdy
Head, NSW, in 990-1020 m.

464 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

veloped. Head lacking stout ridges of modified scales; tip and lateral angles of snout armed with small
tubercular scales with upturned spinules; snout otherwise naked. Body scales large, deciduous; spi-
nules on exposed field reduced or lacking. Long spinous ray of first dorsal fin with few or no spinules
along leading edge. Pectoral fin relatively long, 50-70% HL; rays fine, none thickened or notably pro-
longed beyond others. Outer pelvic ray slightly produced, its length 40-60% HL. Color in alcohol
overall pale to brownish, flesh translucent over anal pterygiophores; blackish over abdomen,
operculum, and somewhat dusky on underside of head; fins pale except for blackish spinous ray of
first dorsal, uppermost ray of pectoral, and outer ray of pelvic.

SIZE. — To about 41 cm.

DISTRIBUTION. — High latitudes of southern hemisphere; known off Australia (NSW, including
Lord Howe Rise, Tas., SA); capture depths range about 1000-5100 m, but most from 3500-5000 m.

NSW CAPTURES. — A single Kapala specimen in 990-1020 m off Crowdy Head and three
Franklin specimens from 2450 m on the Lord Howe Rise.

REMARKS. — Coryphaenoides filicauda is primarily an abyssal species, and the Kapala capture
at mid-slope depths off NSW is the shallowest record for the species. The three juveniles
(17.5—20.1 mm HL) from the Lord Howe Rise differed from others examined (including from other
areas) ina number of features, including slightly more gill rakers on first arch (6—8 outer rakers; | | in-
ner rakers), somewhat longer snout (30-37% HL), wider interorbital (32-29% HL), longer upper jaw
(38-39% HL), and longer barbel (12-13% HL). We are uncertain whether these differences reflect
ontogenetic change or different taxa. A closely related species, C. carapinus, can be distinguished
from C. filicauda by its well-serrated leading edge of the spinous dorsal ray, longer barbel (9-15%
HL), more numerous and better-developed gill rakers (GR-I 6—10/9—13), and more adherent, more
heavily spinulated scales.

REFERENCE SPECIMENS. — AMS I.27643-004 (1 spec.); K88-12-02. AMS 1.29316-004 (3
spec.); Lord Howe Rise, 29°42.06’S, 159°48.13'E; 2450 m; ORV Franklin stn FRO589-17; 3 May
1989.

REFERENCES. — Iwamoto and Sazonov (1988); Gon and Heemstra (1990).

Coryphaenoides grahami lwamoto and Shcherbachey, 1991
Fig. 80

DISTINGUISHING FEATURES. — 1D II,8—9; P i19-i23; V 12; GR-I (total, outer/inner series)
8—10/11—14; scales below 1D 7.5—9.0, below 2D 8.0-9.5, lat.l about 26-31; pyloric caeca about 10.
Snout length 25-29% of HL; orbit diameter 21-23%; interorbital width 28-31%; suborbital width
11-14%; upper jaw length 39-42%; barbel length 13-23%. Head about 6 in TL; preopercle with 4
spikelike struts. Teeth small, weak, somewhat deciduous, in single row in lower jaw. Head lacking

FIGURE 80. Coryphaenoides grahami Iwamoto and Shcherbachev, 1991. AMS 1.29798-010. From Kapala stn K89-12-04,
off Tuncurry, NSW, in 1033-1079 m.

ES OE Ee OM

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 465

stout ridges of large, modified scales and large tubercular scale at snout tip; underside of head (except
gill membranes) entirely scaled; body scales deciduous, about 8—10 parallel rows of small spinules.
Color fresh ivory white with silvery sheen; in alcohol overall dark gray to brown, blackish over ventral
surfaces and lips, mouth, gill membranes, gill chamber, and most fins.

SIZE. — To about 40 cm.

DISTRIBUTION. — South Atlantic off South Africa, southern Indian Ocean, eastern Australia
(NSW), in about 1050-1300 m.

NSW CAPTURES. — Six specimens collected at four Kapala stations between Crowdy Head and
Jervis Bay in 1040-1140 m.

REMARKS. — Coryphaenoides grahami has not been collected from any other Australian state. It
is likely to be mistaken only for C. striaturus, which is similar in shape, and shares many counts, mea-
surements, and scale features. Coryphaenoides grahami is, however, darker overall and has a broader
interorbital, smaller, weaker teeth, and spikelike processes on the preopercle.

REFERENCE SPECIMENS. — AMS _1.29737-002 (paratype), and 1.29737-004 (2. spec.);
K89-19-01. AMS 1I.29742-003 (paratype); K89-17-07. AMS 1.29745-005 (paratype); K89-18-02.
AMS 1.29798-010 (paratype); K89-12-04.

REFERENCE. — Iwamoto and Shcherbachev (1991).

Coryphaenoides mcmillani lwamoto and Shcherbachey, 1991
Fig. 81

DISTINGUISHING FEATURES. — ID II,9—11; P 117-120; V 8—9; GR-I (total, outer/inner series)
11—16/16—19; scales below 1D 7.5—9.0, below 2D 6.5—9.5, lat.1 about 31—34; pyloric caeca about
9-10. Snout length 28-31% of HL; orbit diameter 27-31%; interorbital width 26-30%; suborbital
width 11—12%; upper jaw length 46-50%. Head about 4.5—6 in TL; sensory pores prominent; chin
barbel rudimentary, stumplike. Upper jaw teeth in narrow band, with slightly enlarged outer series;
lower jaw teeth in single row. Outer gill rakers of first arch relatively long and flat, triangular to sa-
ber-shaped. Head lacking stout ridges of large, modified scales; snout naked except for small tubercu-
lar scale at tip and lateral angles; body scales deciduous, about 8—10 parallel rows of slender, conical
spinules. Pectoral fin relatively long, 70-90% HL, but rays fine, none thickened or notably prolonged
beyond others. Outer pelvic ray elongated, its length 70-90% HL. Color in alcohol overall swarthy,
blackish over abdomen behind pelvic fins; operculum and most head membranes black; fins blackish
to dusky.

SIZE. — To about 35 cm.

DISTRIBUTION. — New Zealand, Australia
(NSW, Tas., SA), southern Indian Ocean, south-
ern Africa, South Atlantic off Whale Ridge, in
950-1400 m.

NSW CAPTURES. — Captured once by ORV
Franklin off Nowra in 817-1009 m.

REMARKS. — Coryphaenoides = mcmillani
was not collected by Kapala despite the holotype
coming from an area and depth extensively
trawled. It is either very rare off NSW or was mis-
identified in the field as C. subserrulatus, which
was commonly recorded from the type area. C.
mcemillani is distinguished from the similar C.

subserrulatus by having more pelvic finrays and Figure 81. Coryphaenoides mcmillani Iwamoto and
shorter pectoral and pelvic fins. Shcherbachev, 1991 (from original illustration of holotype).

466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

REFERENCE SPECIMEN: NMV A6794 (holotype); NSW, 56 km off Nowra, 34°44’S, 151°14.3’E,
in 1009-817 m; ORV Franklin stn CSIRO FRS5/86, Slope 9.
REFERENCE. — Iwamoto and Shcherbachev (1991).

Coryphaenoides murrayi Gunther, 1878
Fig. 82

DISTINGUISHING FEATURES. — 1D II,8—10; P 118-120; V 10—12(usually 12); GR-I (total,
outer/inner series) 9—11/14—16; scales below 1D 9-11, below 2D 9-11, lat.1 about 33; pyloric caeca
about 9-10. Snout length 28-30% of HL; orbit diameter 20—23%; interorbital width 28-34%; subor-
bital width 13—17%; upper jaw length 42-45%. Head broad, width about two-thirds its length; chin
barbel slender, about equal to or longer than orbit. Upper jaw teeth in broad band, with outer series
slightly enlarged; lower jaw teeth in one row. Head lacking ridges of large, stout, modified scales; a
row of small scales along leading edge of snout; snout naked on underside, with broad naked areas on
dorsal surface behind leading edge; body scales rather deciduous, about 5—8 parallel rows of weak,
slender, conical spinules. Color in alcohol overall dark brownish to swarthy.

SIZE. — To at least 37 cm.

DISTRIBUTION. — Western Indian Ocean to southeastern Australia (NSW, Vic.), New Zealand,
and Fiji, in depths of 1196-2350 m.

NSW CAPTURE. — Taken once by ORV Franklin off Nowra in 1896-1642 m.

FIGURE 82. Coryphaenoides murrayi Ginther, 1878. Holotype, BMNH 1887.12.7.113, Challenger stn 168, off New Zea-
land, in 2012 m. Fins and scales reconstructed.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 467

REMARKS. — This is a species of lower-slope depths known mostly from captures off New Zea-
land. The only Australian specimens are from off NSW and eastern Victoria.

REFERENCE SPECIMENS. — NMV A7004 (1 spec.); 67 km off Nowra, NSW, 34°41.97'S,
151°22.44'E, in 1896-1642 m; ORV Franklin stn CSIRO FR9/88, Slope 59, 22 Oct 1988. NMV
A6793 (1 spec.); 85 km s. of Pt. Hicks, 38°31.4'S, 149°21.1'E, in 1986-1360 m.

REFERENCE. — Iwamoto and Shcherbachev (1991).

Coryphaenoides rudis Giinther, 1878
Fig. 83

DISTINGUISHING FEATURES. — ID IJ,9—11; P 116-121; V 8-11 (usually 9 or 10, rarely 8 or 11);
GR-II (total, outer/inner series) 7—10/9-10; scales below 1D 9, below 2D 6.5, lat.1 about 37; pyloric
caeca about 14. Head broad, width about two-thirds its length; snout low, scarcely protruding in large
adults, length about 23—29% HL; orbit small, 16-26% HL; interorbital width 26-30% HL; mouth
large, upper jaw extending to below posterior margin of orbit, 37-43% HL; chin barbel slender,
10-23% HL. Upper jaw teeth in moderately broad band, with outer series enlarged; lower jaw teeth in
1-3 irregular rows. Head lacking ridges of large, stout, modified scales; snout completely scaled;
body scales with small spinules. Color in alcohol overall brownish to brownish gray.

SIZE. — One of the largest known grenadiers, attaining at least 120 cm in TL. The two NSW
specimens measured 71 and 95 cm in length.

DISTRIBUTION. — Broadly distributed in central and western Pacific, Indian Ocean, and middle
to low latitudes in the Atlantic, in depths of 1,000—2,400 m. Recorded from Australia off NSW and
WA.

NSW CAPTURES. — Only two specimens caught by Kapala, one in 1050 m off Newcastle, the
second in 1150 m off Sydney.

REMARKS. — Juveniles of this large species have a more protruding snout and proportionately
larger orbits than do the adults, which affects the relative dimensions of the interorbital, suborbital,
and postorbital. Coryphaenoides rudis is a large, widely distributed species originally described from
the Kermadec Islands, but subsequently recorded from other areas under the names C. paradoxus and
C. macrocephalus, as well as C. rudis. It appears to be rare wherever found.

REFERENCE SPECIMENS. — AMS I.28477-001 (1 spec.); K88-16-04; AMS I.29340-001 (1
spec.); K89-13-02.

REFERENCES. — Sazonov and Iwamoto (1992); Shcherbachev and Iwamoto (1995); Iwamoto
and Williams (1999).

Coryphaenoides serrulatus Giinther, 1878
Fig. 84

DISTINGUISHING FEATURES. — 1D II,8—11; P 118-122; V 7 (rarely 6 or 8); GR-I (total, outer/in-
ner series) 7—10/11—15, GR-II 10—14/10—14; scales below 1D 8-10, below 2D 6.5-8.5, lat.1 35-40;
pyloric caeca 16—19. Snout length 27-30% HL; orbit diameter 30-33%; interorbital width 18-24%;
suborbital width 1 1—14%; upper jaw length 39-44%. Head short and compressed, about 6 times in
TL; chin barbel well developed, length 20-30% HL. Upper jaw teeth in relatively narrow band, with
outer series enlarged; lower jaw teeth in one row. Gill rakers rather numerous, the outer rakers some-
what tablike. Suborbital ridge with row of stout, coarsely thickened scales; tip and lateral angles of
snout with large, tubercular scales, snout otherwise uniformly covered with small scales; body scales
rather large, covered with lanceolate spinules. Outer pelvic ray 50-80% HL, extending to anal origin
or slightly beyond. Color in alcohol overall brownish, darker over abdomen and gill covers with blu-
ish to blackish.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

468

Volume 52, No. 21

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IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 469

SIZE. — To about 45 cm.

DISTRIBUTION. — New Zealand and Australia (NSW, Vic., Tas., SA, WA), in 550-1200 m.

NSW CAPTURES. — Caught on all mid-slope grounds south of Crowdy Head in 690-1200 m.
The most abundant grenadier in Kapala mid-slope catches. Recorded from 253 stations and present in
all but 5 trawls in depths between 800 and 1200 m. Average catch about 40 per tow.

REMARKS. — Two subspecies of this widely distributed species of the southern hemisphere were
recognized by Iwamoto and Shcherbachev (1991), with Coryphaenoides s. serrulatus occurring off
New Zealand and southern Australian waters and C. s. oceanus found in oceanic waters of the Indian
Ocean. The latter subspecies differs from the former in having a longer outer pelvic ray that extends
beyond the base of the 10th anal ray, as well as several differences in proportional measurements.

REFERENCE SPECIMENS. — AMS I.18726-025 (3 spec.); K75-01-02. AMS 1.19859-001 (13
spec.); K76-24-04. AMS I.19860-006 (8 spec.); K76-24-03. AMS I.19862-004 (5 spec.); K76-23-01.
AMS 1.20096-007 (8 spec.); K77-22-06. AMS I.20098-006 (3 spec.); K77-23-07. AMS I.20484-002
(1 spec.); K77-21-01. AMS 1.20485-007 (3 spec.); K77-23-06. AMS 1.23885-015 (1. spec.);
K78-27-05. AMS 1.24037-004 (6 spec.); K78-26-16. AMS 1.24054-013 (1 spec.); K83-06-02. AMS
1.24055-009 (1 spec.); K83-08-01. AMS 1I.24172-006 (1 spec.); K83-14-01. AMS 1.24613-001 (8
spec.); K75-05-05. AMS 1.25933-006 (1 spec.); K79-20-06. NMV A17 (1 spec.) and NMV A9077 (2
spec.); 56 km off Nowra; 1009-817 m; ORV Franklin stn CSIRO FRS/86, Slope 9. NMVV A5783 (1
spec.), off Nowra; 1100 m; ORV Franklin stn CSIRO FR9/88, Slope 58.

REFERENCES. — Last et al. (1983); Iwamoto and Shcherbachev (1991); McMillan in Gomon et
al. (1994); Iwamoto and Williams (1999).

Coryphaenoides striaturus Barnard, 1925
Fig. 85

DISTINGUISHING FEATURES. — 1D II,8—10; P 118-124; V 11—12; GR-I (total, outer/inner series)
7—-11/12—16, GR-II 1 1—14/11-—15; scales below 1D 7.5—10, below 2D 8-9, lat.1 30—36; pyloric caeca
9-12. Snout length 26—30% HL; orbit diameter | 8—24%; interorbital width 23—30%; suborbital width
11-15%; upper jaw length 38-46%. Head robust, its width about equal to postorbital length, about 5.5
in TL; chin barbel well developed, length | 8—26% HL. Upper jaw teeth in band, with outer series en-
larged; lower jaw teeth in one row. Gill rakers somewhat tablike. Head ridges prominent but not rein-
forced by thickened scales; underside of snout covered with small scales; body scales rather large,
with 9-11 parallel rows of small, needlelike spinules. Outer pelvic ray extends beyond anal fin origin,

rs

a

FiGuRE 85. Coryphaenoides striaturus Barnard, 1925. AMS 1.29737-007. From Kapala stn K89-19-01, off Ulladulla, NSW,
in 1116-1134 m.

470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

usually to base of 6th—1 Ith ray. Color in alcohol overall dark brown to swarthy, darker on underside of
head and gill covers.

SIZE. — To about 55 cm.

DISTRIBUTION. — New Zealand, Australia (Qld, NSW, Vic., Tas., SA, WA), to the southeastern
Atlantic off southern Africa, in depths of about 800 to 2000 m; most often taken in 1000—1400 m.
Generally confined to cooler, temperate waters off southern coasts, although two AMS specimens
were taken on the Lord Howe Rise off southern Qld and northern NSW.

NSW CAPTURES. — Collected from only three Kapala stations (5 specimens) south of 34°50'S
in about 1100 m depth. A fourth NSW specimen was collected by ORV Franklin on the Lord Howe
Rise in 1590 m.

REMARKS. — Coryphaenoides striaturus is normally an abundant species throughout its range,
but off NSW it may be more common in depths greater than those trawled by Kapala. The uniformly
and completely scaled snout, the pelvic fin ray count, and the highly arched nape help to distinguish
this species from other Australian members of the subgenus Chalinura. The sympatric species C.
grahami is closely similar but can be differentiated by its spikelike ridges of the preopercle, darker
overall color, and less arched nape. Also, C. striaturus is a stouter, firmer-fleshed fish than is C.
grahami.

REFERENCE SPECIMENS. — AMS 1.24992-004 (2 spec.); K84-11-07. AMS 1.29737-007 (2
spec.); K89-19-01. AMS 1.29745-002 (1 spec.) and AMS 1.29745-003 (1 spec.); K89-18-02. AMS
1.293 15-002 (1 spec.); Lord Howe Rise, 29°10.29’S, 160°29.78'E; 1590 m; 4 May 1989; ORV Frank-
lin stn FRO589-21. AMS 1.29339-001 (1 spec.); Lord Howe Rise, 27°39.8'S, 161°46.38'E; 1423 m;
1989; ORV Franklin stn FRO589-3 1.

REFERENCES. — Iwamoto and Shcherbachev (1991); Iwamoto and Williams (1999).

Coryphaenoides subserrulatus Makushok, 1976
Fig. 86

DISTINGUISHING FEATURES. — 1D II,9-11; P 113-118; V 7; GR-I (total, outer/inner se-
ries) 10—14/16—19, GR-II 16—18/14—17; scales below 1D 8-9, below 2D 6.5-8.5, lat.1 29-37; pyloric
caeca 12—14. Snout length 25-29% HL; orbit diameter 30-34%; interorbital width 20-25%; subor-
bital width 7—10%; upper jaw length 44-49%. Head short and laterally compressed, more than 6 times

FIGURE 86. Coryphaenoides subserrulatus Makushok, 1967. AMS 1.24054-006. From Kapala stn K83-06-02, off
Wollongong, NSW, in 869-878 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 47]

in TL; chin barbel rudimentary. Upper jaw teeth in relatively narrow band, with outer series slightly
enlarged; lower jaw teeth in one row. Gill rakers numerous for genus, the outer rakers on first arch
somewhat tablike. Suborbital shelf narrow, with row of stout, coarsely thickened scales; tip and lateral
angles of snout with large, tubercular scales, snout otherwise mostly naked ventrally and partly dor-
sally; body scales rather large, covered with lanceolate spinules. Uppermost developed pectoral fin
ray stouter than other rays of fin and greatly elongated, 115—214% HL. Outer pelvic ray elongated,
158—221% HL, extending far beyond anal fin origin. Color in alcohol overall light to medium brown,
darker over abdomen, gill covers bluish to blackish.

SIZE. — To 37 cm.

DISTRIBUTION. — Widely distributed off southeastern Australia (NSW, Vic., Tas.), New Zea-
land, Chile, and in the South Atlantic off Argentina and on the Agulhas Plateau. Depth range about
700-1200 m, but one capture off Chile at 470-440 m.

NSW CAPTURES. — Commonly caught on all grounds south of Crowdy Head in 720—1200 m.
Recorded from 173 Kapala stations including 70% of stations deeper than 800 m; average catch about
10 per trawl.

REMARKS. — Coryphaenoides subserrulatus differs from the closely similar C. memillani in the
relatively thick, greatly elongated uppermost pectoral fin ray (no prolonged ray in C. mcmillani),
fewer pelvic fin rays (7 cf. 8 or 9), and lanceolate scale spinules (cf. needlelike, in parallel rows).

REFERENCE SPECIMENS. — AMS 1I.20068-005 (1 spec.); K77-23-13. AMS 1.24037-003 (7
spec.); K78-26-16. AMS I.24054-006 (5 spec.); K83-06-02. AMS I.24055-004 (1 spec); K83-08-01.

REFERENCES. — Makushok (1967); McCann and McKnight (1980) (as Coryphaenoides
quadripennatus); Iwamoto and Shcherbachev (1991).

Genus Cynomacrurus

DISTINGUISHING FEATURES. — Branchiostegal rays 6. Mouth large, upper jaw extending well
past orbit. Anus located immediately before anal fin origin; no light organ. Swim bladder very small.
Sensory pores large, prominent; orbit small, more than 5 in head length. One or more pairs of large
fanglike teeth in upper jaw; lower jaw with single row of 4 to 11 fanglike teeth. Lateral line broken into
two main segments, anterodorsal segment ending somewhat behind first dorsal fin, second segment
midlateral, beginning below end of first segment.

REMARKS. — Monotypic genus; bathypelagic in Southern Ocean.

REFERENCES. — Dollo (1909); Marshall (1964); Iwamoto in Gon and Heemstra (1990).

Cynomacrurus piriei Dollo, 1909
Fig. 87

DISTINGUISHING FEATURES. — ID II,8—-9; P 113-116; V 7-8; GR-I (outer/inner) 8 or 9 to-
tal/14—16 total; scale rows below 1D about 7; pyloric caeca 9 or 10. Head about 5 times in total length;
snout not protruding. Extensive naked areas on head. Barbel absent. Spinous second ray of first dorsal
fin smooth.

SIZE. — To about 46 cm.

DISTRIBUTION. — Southern Ocean, Australia (NSW), and New Zealand (fide Peter McMillan).

NSW CAPTURE: One specimen (the only Australian record) captured by Kapala off Ulladulla
(35°30'S) when trawling in 1030-1070 m. It may have been captured in midwater when hauling the
trawl.

REMARKS. — The species is abundant at bathypelagic depths of the Southern Ocean.

REFERENCE SPECIMEN: AMS 1I.24424-005 (1 spec.); K83-19-02.

472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 87. Cynomacrurus piriei Dollo, 1900. AMS 1.24424-005. From Kapala stn K83-19-02, off Ulladulla, NSW, in
1079-1116 m.

Genus Haplomacrourus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Head laterally compressed, snout
rounded in profile; in larger specimens (>30 cm TL) upper jaws fall short of vertical through anterior
margin of orbit and snout not protruding beyond mouth. Snout and ventral parts of head naked; scales
small, those on head and anterior part of body lacking spinules. Anus about midway between anal and
pelvic fins; a small dermal window of light organ between bases of pelvic fins. Spinous dorsal ray
stout, flattened laterally, recurved and finely serrated along leading edge.

REMARKS. — A peculiar monotypic species of uncertain relationships.

REFERENCES. — Trunov (1980); Iwamoto and Merrett (1997).

Haplomacrourus nudirostris Trunovy, 1980
Fig. 88

DISTINGUISHING FEATURES. — 1D II,9—10; P i25-i28; V 8—9; scales small, 15-18 below ID,
15—17 below 2D; pyloric caeca about 30.
SIZE. — To about 60 cm.

FIGURE 88. Haplomacrourus nudirostris Trunov, 1980. AMS 1.26247-004. From Kapala stn K85-21-04, east of Broken
Bay, NSW, in 1024-1052 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 473

DISTRIBUTION. — Southeastern Atlantic off Africa through Indian Ocean to Australia (NSW)
and New Zealand, north to New Caledonia, in about 800—1600 m.

NSW CAPTURES. — Caught at eight stations (9 specimens) between Crowdy Head and Jervis
Bay in 950-1100 m.

REMARKS. — All Kapala specimens were small (174-280 mm TL), overall bluish black (trunk
blackish), with a rounded snout profile, and jaws extending posterior to a vertical through the anterior
margin of the orbit. The small numbers and absence of large adults in Kapala catches suggests that
Haplomacrourus more commonly inhabits depths greater than those fished, or the adults are avoiding
the net, or that NSW is marginal to its normal distribution.

REFERENCE SPECIMENS. — AMS 1.24057-005 (2 spec.); K83-09-04. AMS 1.24993-001 (1
spec.); K84-16-14. AMS I.26247-004 (1 spec.); K85-21-04. AMS I.29752-001 (1 spec.); K89-17-08.
AMS 1.29754-004 (1 spec.); K89-17-03. AMS 1I.29797-002 (1 spec.); K89-12-05. AMS 1.29799-001
(1 spec.); K89-06-02. AMS 1.40272-004 (1 spec.); K80-20-05.

Genus Hymenocephalus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Head bones weakly ossified, head cover-
ing membranous, transparent. Spinous ray of first dorsal fin usually smooth (weakly serrated in sub-
genus Hymenogadus, species of which may occur off NSW but have yet to be recorded). Anus
immediately before anal fin origin, without broad black naked perianal margin. Two lens-like struc-
tures of light organ, one on chest, the other immediately before anus; luminescent tissue, consisting of
fine black striations between silver ground, cover parts of abdomen, chest, shoulder girdle and isth-
mus between gill membranes. Gill rakers tubercular, inner rakers of first arch usually more than 18.
Most species small, usually < 20 cm TL. Color blackish, with silvery cover over most of head and
ventral surfaces of body; but some species almost entirely black.

REMARKS. — Only three species of this widespread genus were identified from Kapala catches,
but others could be expected off northern NSW, especially those species reported from New Caledo-
nia by Iwamoto and Merrett (1997).

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto and Merrett (1997).

KEY TO THE SPECIES OF HYMENOCEPHALUS IN NEW SOUTH WALES

la. Chin barbel long, well developed; pelvic finrays8...............0 002005 e eee eee H. longibarbis
1b. Chin barbel absent or rudimentary; pelvic finrays11-14................. 2.00.00 20 20552 e eee 2
2a. Orbits small, 3.6—4.5 times into HL; midlateral dark stripe faint or inconspicuous; body rather uniformly dark,
fade posteriorly; suborbital broad, 10-16 into. orbit. oc). os cee eo wk ee ee we H. aterrimus
2b. Orbits large, 3.2 or less times in HL; distinct midlateral dark stripe present extending to end of tail; suborbital 2-3
HLILQIOTOIME oe clades ea ee CSP EMA She ee, ES TR Re ens Qe cee SID ae, EM ees H. nascens

Hymenocephalus aterrimus Gilbert, 1905
Fig. 89

DISTINGUISHING FEATURES. — V 12-14; total GR-I (outer/inner) |4—20/22—27. Chin barbel ab-
sent. Suborbital region broad, width 17—22% HL; interorbital broad, width 36-39% HL; orbit small,
diameter 22—28% HL. Color uniformly black to dark brown in preserved specimens, generally paler
on tail.

SIZE. — To about 19 cm.

DISTRIBUTION. — Widespread in warm waters of Indian, Atlantic, and Pacific oceans, including
Australia (NSW). Questionably recorded from the southeastern Pacific off Sala-y-Gomez and Nazca
ridges (Sazonov and Iwamoto 1992), but otherwise not present in tropical eastern Pacific.

474 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 89. Hymenocephalus aterrimus Gilbert, 1905. AMS 1.29753-001. From Kapala stn K89-17-06, east of Crowdy
Head, NSW, in 878-933 m.

NSW CAPTURES. — Collected twice in 820-830 m off Crowdy Head at the northern end of the
mid-slope trawling ground.

REMARKS. — The NSW specimens are the only Australian records, but H. aterrimus can be ex-
pected off Queensland. Its small size and more tropical distribution could account for its rarity in
Kapala trawls. This species deserves closer scrutiny, as specimens recorded from widely separated
areas under the name may represent more than one species.

REFERENCE SPECIMENS. — AMS 1.29742-001 (1 spec.); K89-17-09. AMS 1.30304-006 (3
spec.); K89-17-06.

REFERENCES. — Gilbert (1905); Sazonov and Iwamoto (1992); Iwamoto and Merrett (1997).

Hymenocephalus longibarbis (Ginther, 1887)
Fig. 90

DISTINGUISHING FEATURES. — V 8; total GR-I (outer/inner) 12—16/19—22. Chin barbel well de-
veloped, its length 38-58% HL. Body long and slender, head rather shallow, suborbital region nar-
row, its width 8-10% HL, interorbital width 16-23% HL, orbit large, diameter 32-41% HL. Color
when fresh overall silvery, with grayish dorsally and blackish ventrally on trunk and over gill mem-
branes; in alcohol silvery color often lost, lateral stripe somewhat diffuse in larger specimens but
prominent on trunk in smaller specimens. Ventral surfaces of tail lacking pigmentation.

DISTRIBUTION. — Fiji, New Caledonia region, Australia (Qld, NSW, possibly WA), and possi-
bly Indonesia.

NSW CAPTURES. — Collected by Kapala between southern Queensland (27°50'S) and
Ulladulla (35°30'S) in 360-820 m. Recently collected in small numbers southeast of Bermagui
(36°30'S)(AMS 1.40289-002). Recorded from 102 Kapala stations with its greatest abundance in
400-600 m off central and northern NSW. Because of its small size, few were collected in fish trawls
fitted with 90 mm mesh codends (caught in only 6% of fish trawls in 400-600 m). In contrast, H.
longibarbis was recorded from 62% of prawn trawl stations in 400-600 m (45 mm mesh nets). Tows
with prawn trawls frequently captured 25—S0 specimens.

TWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 475

FIGURE 90. Hymenocephalus longibarbis Ginther, 1877. From Kapala stn K83-01-08 off Sydney, NSW, in 490-570 m.

REMARKS. — Hymenocephalus longibarbis is abundant off Queensland, and Iwamoto and Wil-
liams (1999) also recorded the species from two captures on the North West Shelf off WA. Their spec-
imens had notably different proportional measurements of the orbit diameter, interorbital width, and
suborbital width, which led them to suggest that they might represent another species.

REFERENCE SPECIMENS. — AMS I.20071-041 (1 spec.); K77-19-05. AMS 1.20118-034 (11
spec.); K77-13-10. AMS 1.20301-024 (3 spec.); K77-13-12. AMS 1I.20518-005 (4 spec.), AMS
1.20518-013 (1 spec.); K78-09-05. AMS 1I.21669-003 (3 spec.); K77-16-16. AMS 1.21795-006 (3
spec.); K78-23-08. AMS I.21805-001 (2 spec.); K77-23-09. AMS I.21806-003 (1 spec.); K77-07-10.
AMS 1.23486-001 (1 spec.); K82-24-02. AMS I.24619-006 (1 spec.); K81-17-03. AMS I.24850-001
(3 spec.); K84-15-01. AMS 1.24852-010 (1 spec.); K84-15-03. AMS 1.29535-002 (7 spec.);
K79-15-01. AMS 1.30407-004 (3 spec.); K78-01-01. AMS 1.40289-002 (3 spec.); FV Shelley H;
36°30'S, 150°21'E; 390-558 m; off Bermagui, NSW; 18 April 2000. AMS 1.40292-002 (3 spec.); FV
Shelley H; 36°26'S, 150°21'E; 428-468 m; off Bermagui, NSW; 2 May 2000.

REFERENCES. — Paxton et al. (1989)(as H. longiceps, in part); Iwamoto and Merrett (1997);
Iwamoto and Williams (1999).

Hymenocephalus nascens Gilbert and Hubbs, 1920
Fig. 91

DISTINGUISHING FEATURES. —V 11-13, usually 11 or 12; total GR-I (outer/inner)
16—20/22-27. Chin barbel absent. Snout conically pointed in lateral view, projecting well beyond
mouth. Suborbital region narrow, width 12-16% HL, interorbital width 1.0—1.4 into orbit, width
27-35% HL, orbit large, diameter 31-38% HL. Color mostly silvery ventrally on head and body; in
preservative, a prominent dark lateral stripe present (silvery when fresh). (After Iwamoto and Merrett,
1997.)

SIZE. — To about 16 cm.

DISTRIBUTION. — Widespread in tropical western Pacific through Philippines, Indonesia, and
northern Australia (Qld, NSW, WA) in depths of about 350-800 m.

NSW CAPTURES. — Captured only twice by Kapala, in 820-930 m off Crowdy Head (at same
stations as H. aterrimus).

476 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 91. Hymenocephalus nascens Gilbert and Hubbs, 1920. AMS 1.29753-012. From Kapala stn K89-17-06, east of
Crowdy Head, NSW, in 878-933 m.

REMARKS. — As with H. aterrimus, this species is marginal to NSW, having its main distribution
in warmer tropical waters; it is abundant in the New Caledonian region. Its small size also made it un-
likely to be captured in Kapala’s trawls.

REFERENCE SPECIMENS. — AMS 1I.29753-012 (1 spec.); K89-17-06. AMS 1.30304-005 (1
spec.); K89-17-09.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto and Merrett (1997); Iwamoto and Wil-
liams (1999).

Genus Kuronezumia

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Anus situated in an oval to tear-
drop-shaped (periproct) area between pelvic fin base and anal fin origin, usually closer to former; a
small shallow black pit representing dermal window of light organ anterior to periproct and between
pelvic fin bases. Body deep, laterally compressed, depth 90—1 10% HL. Head usually much deeper
than wide; almost completely and uniformly covered with small spinulated scales, including those
over broad, flat suborbital region; snout somewhat rounded or bluntly protruding; mouth moderate in
size, upper jaw 30-44% HL. Second spinous ray of first dorsal fin serrated along leading edge. Teeth
in broad bands in both jaws. Gill rakers on outer side of second arch 8—11 total. Color light gray to
brown to swarthy, but lacking bluish or violet.

REMARKS. — Five species of this genus are currently recognized, with two represented in NSW.
Members of the genus are similar to some species of Nezumia, especially in terms of their overall
physiognomy, but they can be distinguished by their almost entirely scaled head and the absence of a
well-developed double row of enlarged, thickened scales along the suborbital ridge.

REFERENCES. — Iwamoto (1974); Shcherbachev et al. (1992)

KEY TO THE SPECIES OF KURONEZUMIA IN NEW SOUTH WALES

la. Pelvic fin rays 11-13; snout rounded, not terminating in a large tubercular scale; upper jaw length 35-44% HL;

scales rows between origin of second dorsal fin and lateral line 12-15... .............-.2-. K. bubonis
1b. Pelvic fin rays 8-9, rarely 10; snout bluntly pointed, terminating in a large tubercular scale; upper jaw length
28-35% HL; scales between origin of second dorsal fin and lateral line 11-12. ................ K. leonis

Kuronezumia bubonis (Iwamoto, 1974)
Fig. 92

DISTINGUISHING FEATURES. — D II,10—-12; P i21-125; V 11-13; total GR-I (outer/inner)
6—8/8—1 1 total; scale rows below | D about 14—21; pyloric caeca 35—39. Snout length 26-32% of HL,
orbit diameter 23-31%, interorbital width 23-26%, height of first dorsal fin about 90%. Snout

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 477

FIGURE 92. Kuronezumia bubonis (Iwamoto, 1974). AMS 1.24645-004. From Kapala stn K84-06-03, off Broken Bay,
NSW, in 777-823 m.

rounded, scarcely protruding beyond mouth, not tipped with an enlarged spiny scute. A large, scaly,
tubercular swelling between pelvic fin bases.

SIZE. — To more than 73 cm.

DISTRIBUTION. — Known from the western Atlantic, Hawaii, South China Sea, southern Indian
Ocean, New Zealand, and eastern Australia (NSW) in depths of around 600 to about 1100 m.

NSW CAPTURES. — Uncommon; 31 specimens caught at 23 Kapala stations on grounds be-
tween Crowdy Head and Jervis Bay in 670-1010 m. Within its main depth range (700-900 m), K.
bubonis was recorded in 21% of trawls.

REMARKS. — So far only reported in Australian waters from NSW; the record by Paxton et al.
(1989) off Cape Everard, Vic., was a misidentification of K. /eonis. It is somewhat peculiar that the
only Kuronezumia collected off Western Australia by Iwamoto and Williams (1999) was a related
species, K. pallida (Sazonov and Iwamoto, 1992), a species previously known only from the south-
eastern Pacific. The Kuronezumia specimens recorded from the Indian Ocean by Shcherbachev
(1987) should be re-examined in this light.

REFERENCE SPECIMENS. — AMS I.17316-008 (1 spec.); K72-07-15. AMS 1.17859-002 (2
spec.); K72-06-06. AMS 1.17867-008 (2 spec.); K72-07-04. AMS 1I.18726-020 (1 spec.), AMS
1.18726-021 (i spec.); K75-01-02. AMS I.19860-015 (1 spec.); K76-24-03. AMS 1I.21722-001 (1
spec.); K79-20-13. AMS I.24054-001 (2 spec.); K83-06-02. AMS I.24101-010 (1 spec.); K83-06-01.
AMS 1.24645-004 (1 spec.); K84-06-03. AMS 1.24991-002 (2 spec.); K84-16-05.

REFERENCES. — Iwamoto (1974); Shcherbachev et al. (1992).

Kuronezumia leonis (Barnard, 1925)
Fig. 93

DISTINGUISHING FEATURES. —D II,9-10; P il9-i26; V 8-10; total GR-I (outer/inner)
8—11/7—12; scale rows below 1D 17-20; pyloric caeca 14—18. Snout length 27-34% HL, orbit diame-
ter 27-35%, interorbital width 22-30%, height first dorsal fin about 70-90%. Snout bluntly pointed,
tipped with an enlarged, buttonlike spiny scute.

SIZE. — To about 50 cm.

478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 93. Kuronezumia leonis (Barnard, 1925). AMS 1.28071-001. From Kapala stn K88-04-09, off Broken Bay, NSW, in
905-969 m.

DISTRIBUTION. — South Atlantic, southern Indian Ocean, Australia (NSW, Vic., Tas., WA), and
New Zealand, in depths of around 700—1100 m.

NSW CAPTURES. — Caught on all mid-slope grounds south of Crowdy Head in 730-1180 m.
The species was common but never taken in large numbers; it was recorded from 147 Kapala stations
(61% of trawls deeper than 800 m) with an average of four specimens per tow.

REMARKS. — Kuronezumia leonis was taken on three occasions with its congener K. bubonis,
but that species has a generally shallower depth range and is much less common.

REFERENCE SPECIMENS. — AMS I.18726-017 (1 spec.); K75-01-02. AMS I. 19860-010 (1
spec.); K76-24-03. AMS I.20068-01 1 (4 spec.); K77-23-13. AMS I.20096-008 (1 spec.); K77-22-06.
AMS 1I.20098-005 (4 spec.), AMS 1I.20098-017 (1 spec.), AMS 1.20098-024 (1 spec.); K77-23-07.
AMS 1.20099-008 (1 spec.), AMS I.20099-019 (2 spec.); K77-23-12. AMS 1.24054-005 (3 spec.),
AMS 1.24054-011 (1 spec.), AMS 1.24054-018 (1 spec.); K83-06-02. AMS I.24056-002 (2 spec.);
K83-08-02. AMS 1I.24060-013 (2 spec.); K83-09-01. AMS I.24157-003 (1 spec.); K83-12-04. AMS
1.24356-004 (1 spec.), AMS 1.24356-006 (1 spec.); K83-14-05. AMS 1.24462-004 (1 spec.);
K83-15-02. AMS I.25933-001 (2 spec.), AMS I.25933-003 (4 spec.); K79-20-06. AMS 1.27637-003
(1 spec.); K88-14-04. AMS I.28071-001 (1 spec.); K88-04-09.

REFERENCES. — Iwamoto (1986); Shcherbachev et al. (1992); Iwamoto and Williams (1999).

Genus Lepidorhynchus

DISTINGUISHING FEATURES. — Branchiostegal rays 6. Anus immediately before anal fin. Head
and body laterally compressed, much deeper than wide; head covering thin, somewhat transparent.
Snout scarcely or not protruding. Mouth large, premaxillary extends to hind one-third of orbit. Small
teeth in narrow band in upper jaw with outer row of widely spaced canines; lower jaw teeth in one row,
teeth larger laterally. Scales thin, deciduous, covered with short needlelike spinules, none thickened
or enlarged. Spinous dorsal fin ray smooth along leading edge. Light-producing tissue appearing as
fine black striations cover broad areas ventrally on trunk forward to isthmus and dorsally onto pectoral
girdle to pectoral base, over abdomen, and above anterior part of anal fin; a small naked fossa of light
organ adjacent to anus. Color overall silvery; dorsum grayish green; gill membranes, lower jaw, and
abdomen black.

REMARKS. — Monotypic. Relationships of the genus are obscure, but there are some resem-
blances to Hymenocephalus, especially in regards to the nature of the luminescent tissue on the body.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 479

REFERENCES. — McCulloch (1926); McCann and McKnight (1980); Gomon et al. (1994).

Lepidorhynchus denticulatus Richardson, 1846
Fig. 94

DISTINGUISHING FEATURES. — 1D II,10—11; P 116-118; V 8-9; total GR-I (outer/inner) about
9/16—19. Orbit large, diameter about 30-40% HL, much greater than interorbital space. Chin barbel
small.

SIZE. — To about 55 cm.

DISTRIBUTION. — Southern Australia (NSW, Vic., Tas., SA, WA), New Zealand, Kermadec Is-
lands, in depths less than 100 to more than 1000 m, but most frequent at 300—700 m.

NSW CAPTURES. — Probably the most abundant grenadier off NSW. Captured by Kapala in 603
trawls on all slope grounds south of the Clarence River (29°40’S) in 230-1080 m, the greatest depth
range recorded for any NSW grenadier (apart from some bathyal species). It was most abundant be-
tween 300 and 900 m, where it was captured at over 70% of stations. In the 1996—97 survey using nets
with 90 mm codend mesh, the species was caught in 117 of 165 tows between 220 and 630 m, and the
mean catch was 164 per one-hour tow.

REMARKS. — This species is the most common grenadier off southeastern Australia. Juveniles
are found mostly between 200 and 600 m, and adults usually in depths greater than 500 m. Last et al.
(1983) reported that off Tasmania it is “frequently caught by the tonne as a large part of the bycatch of
trawlers,” and it “appears to be an important prey item for the economically important blue grenadier
[Macruronus novaezelandiae].” Although of no market value at present, the species is frequently
used as tuna longline bait.

REFERENCE SPECIMENS. — AMS 1I.15969-001 (2 spec.); K71-05-06. AMS 1.18838-008 (14
spec.); K75-05-03. AMS I. 21724-021 (2 spec.); K79-20-15. AMS 1.24619-009 (4 spec.); K88-17-03.

REFERENCES. — McCann and McKnight (1980); McMillan in Gomon et al. (1994).

Genus Lucigadus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Anus removed from anal fin, closer to
pelvic fin bases. Light organ well developed, two dermal windows, one immediately before anus, the
second between bases of pelvic fins. Head smoothly rounded, without sharp or coarsely scaled ridges;
snout rounded. Ventral region of body appearing to have swung far forward so that gill membranes

FiGurE 94. Lepidorhynchus denticulatus Richardson, 1846. From Kapala stn K83-01-08 off Sydney, NSW, in 490-570 m.

480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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unite below orbits, pelvic fin origin below opercle, anal fin origin under first dorsal fin. Suborbital
shelf covered with several rows of small scales, no sharp ridge of modified scales. Underside of snout
all or mostly scaled. Spinous ray of first dorsal fin serrated along leading edge. Teeth in both jaws
small, in tapered bands; premaxillary band not reaching beyond posterior edge of maxillary process.
Scale spinules aligned in parallel rows. (From Iwamoto and Merrett 1997.)

REMARKS. — This genus is close to Ventrifossa and Malacocephalus and was formerly included
in the former as a subgenus. The high, bluntly rounded snout, the short trunk with only 10 or 11 trunk
vertebrae, the forward position of the ventral parts of the trunk, the shorter band of premaxillary teeth,
the characteristic arrangement of spinule rows on the scales, and the frequent presence of scales on the —
branchiostegal membrane, in combination distinguish members of the genus from those of
Ventrifossa. Malacocephalus species are readily distinguished by the longer jaws beset with larger
teeth in fewer, longer rows.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto and Merrett (1997).

Lucigadus microlepis (Ginther, 1878)
Fig. 95

DISTINGUISHING FEATURES. — 1D II,10—12; P 120-125; V 11-13; total GR-I (outer/inner)
8—-10/11—12; scales below 2D 8.5—10.5. Snout length 24-31% HL, interorbital width 25-33%, orbit
diameter 32-42%; suborbital width 12—16%; upper jaw length 34-40%; barbel length 18-30%;
height 1D 102—128. A prominent black blotch at tip of first dorsal fin; body with banded pattern, the
anal fin with distinct black margins under darkly banded areas; few or no scales on branchiostegal and
gular membranes.

SIZE. — To about 20 cm.

DISTRIBUTION. — Western South Pacific from Wallis and Futuna islands, Fiji, New Caledonia,
Norfolk Ridge, eastern Australia (Qld, NSW), and Arafura and Madura seas. Depth range about
200-700 m.

NSW CAPTURES. — The Kapala captured the species at 14 stations (36 specimens) on grounds
north of Sydney in the relatively narrow depth range of 410 to 540 m. The small adult size of the spe-
cies made capture in large-meshed trawls unlikely, and in fact, all records were from stations using
prawn trawls. The species was taken from 17% of prawn trawls north of Sydney at 400—600 m depth.

y

Y)

FiGURE 95. Lucigadus microlepis (Ginther, 1878). AMS 1.25932-007. From Kapala stn K85-21-06, off Sydney, NSW, in
439-466 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 481

The overall low capture rate by Kapala suggests that NSW is marginal to its normal tropical distribu-
tion.

REMARKS. — Lucigadus microlepis is readily distinguished from L. nigromaculatus by its
banded body pattern, the head peppered with large melanophores, the location of the black dorsal fin
blotch (to distal tip, compared to below distal tip in L. nigromaculatus), and its somewhat fewer pelvic
fin rays (11-13 cf. 13-15).

REFERENCE SPECIMENS. — AMS 1.20435-015 (1 spec.); K78-16-07. AMS 1.21725-003 (1
spec.); K80-05-01. AMS I. 23689-002 (1 spec.); K78-17-07. AMS I.24850-002 (7 spec.); K84-15-01.
AMS 1I.25932-007 (1 spec.); K85-21-06. AMS I.26394-001 (2 spec.); K86-01-05. AMS 1.26446-008
(2 spec.); K85-17-02. AMS 1.26453-004 (1 spec.); K86-10-07. AMS 1.26756-005 (1 spec.);
K86-10-14. AMS I.26932-004 (1 spec.); K79-15-03.

REFERENCES. — Paxton et al. (1989)(as Ventrifossa fasciata); Iwamoto and Merrett (1997);
Iwamoto and Williams (1999).

Lucigadus nigromaculatus (McCulloch, 1907)
Fig. 96

DISTINGUISHING FEATURES. — ID II,10—11; P il8-i22; V 13-15; total GR-I (outer/inner)
9-1] 1/12—16; scales below 2D 10-12. Snout length 25-30% HL, interorbital width 20-26%, orbit di-
ameter 40-47%; suborbital width 12—18%, upper jaw length 39-45%; barbel length 18-26%; height
1D 101—134%. A prominent black blotch across anterior half to two-thirds of first dorsal fin; anterior
margin of anal fin blackish. Scale patches present on lowermost branchiostegal rays.

SIZE. — To about 35 cm.

DISTRIBUTION. — Southeastern Australia (southern Qld, NSW, Vic., Tas.), New Zealand, and
off Chile. Depth range about 200-1460 m, but most often taken at 400-800 m.

NSW CAPTURES. — Recorded from 222 Kapala stations along the entire NSW coast in depths
between 380 and 850 m. In the 1996-97 survey, L. nigromaculatus was caught in 70% of tows in
440-630 m, with a mean catch of 15 per one-hour tow; most trawls took less than 20 specimens, but
150 were caught in a single tow in 500 m off Ulladulla.

REMARKS. — This species is captured frequently throughout its normal distribution range but
seldom in great abundance. Mesh size may be a factor in its reported low abundance in commercial
trawls, as the species is relatively small as an adult.

FiGuRE 96. Lucigadus nigromaculatus McCulloch, 1907. From Kapala stn K84- 18-03, off Nowra, NSW, in 732-750 m.

482 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

REFERENCE SPECIMENS. — AMS I.15974-008 (4 spec.); K71-08-03. AMS I.15975-032 (4
spec.); K71-08-05. AMS I.15987-001 (4 spec.); K71-11-09. AMS I.15995-011 (2 spec.); K71-13-06.
AMS I.16577-009 (1 spec.); K72-04-02. AMS I.18838-035 (1 spec.); K75-05-03. AMS I.18839-005
(23 spec.), AMS I.18839-012 (4 spec.); K75-05-04. AMS 1.19076-003 (1 spec.); K75-05-08. AMS
1.19198-003 (3 spec.); K76-05-04. AMS 1.20118-003 (2 spec.); K77-13-10. AMS 1.20484-004 (3
spec.); K77-21-01. AMS I.21669-007 (1 spec.); K77-16-16. AMS 1.21806-005 (2 spec.); K77-07-10.
AMS 1.2485 1-004 (1 spec.); K84-17-03. AMS I.24852-003 (1 spec.); K84-15-03. AMS I.26002-001
(1 spec.); K86-01-06. AMS 1.26245-013 (1 spec.); K86-01-07.

REFERENCES. — Paxton et al. (1989)(as Ventrifossa nigromaculata); McMillan in Gomon et al.
(1994).

Genus Malacocephalus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Anus removed from anal fin, closer to
pelvic fin bases. Head laterally compressed; snout rounded, without coarse, thickened scales. Head
surfaces uniformly scaled, no sharp ridge of modified scales, no naked areas; branchiostegal rays
scaled. Spinous ray of first dorsal fin smooth or serrated along leading edge. Light organ well devel-
oped, two dermal windows, one immediately before anus, the second (large and somewhat
bean-shaped) between bases of pelvic fins. Teeth in upper jaw in two rows to narrow band, outer series
enlarged; lower jaw with a single row of wide-spaced, enlarged, canine-like teeth. Scales of body
densely covered with small fine scales giving velvety surface. Pyloric caeca numerous, 50—100. Color
light gray to swarthy, often with silvery flanks.

REMARKS. — There are seven named species, but the number of valid species is uncertain. Three
Pacific species (M. hawaiiensis, M. luzonensis, M. nipponensis) are so closely similar to M. laevis that
the four may eventually prove to be the same, in which case M. /aevis has priority as the oldest name.

REFERENCES. — Iwamoto (1990); Sazonov and Iwamoto (1992).

Malacocephalus laevis (Lowe, 1843)
Fig. 97

DISTINGUISHING FEATURES. — 1D II,9-13; P i15—i21 (usually 117-119); V 9; total GR-I
(outer/inner) 0-8—11/11—14; scales below 2D 8.5—11.5. Snout length 26-31% HL, interorbital width
28-33%, orbit diameter 30-37%; suborbital width 10-13%, upper jaw length 44-50%; barbel length

FIGURE 97. Malacocephalus laevis (Lowe, 1843). From Kapala stn K83-01-08, off Sydney, NSW, in 490-570 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 483

16-23%; height 1D about 60-70%. Two rows of teeth in upper jaw. Spinous ray of first dorsal fin
smooth.

SIZE. — To more than 65 cm.

DISTRIBUTION. — Widespread in tropical to temperate waters throughout Atlantic and Indian
oceans, and probably into Pacific (but greatly restricted in eastern Pacific). Depths approximately
200—1000 m, but most commonly in about 300-700 m. Known off Australia (Qld, NSW, ne. Vic.,
WA).

NSW CAPTURES. — Captured by Kapala on all grounds between the Qld-NSW border and
Batemans Bay (35°42'S); single specimens were also caught in three trawls off ne. Vic.
(37°37'-38°02'S). All stations but two were in 330-800 m; the two other records were in
1030-1080 m. Regularly taken in moderate numbers on the upper slope off NSW. Malacocephalus
laevis was recorded 290 times by the Kapala; north of Jervis Bay, it was present in 78% of all trawls in
400—800m. Overall, the average catch was about 10 specimens per trawl, although some tows caught
in excess of 100.

REMARKS. — Malacocephalus laevis is a well-known, apparently worldwide species with a
mainly tropical and subtropical distribution. It appears to be found only in relatively warm waters
around Australia. Off NSW few were caught south of 35°00'S, and it appears to be absent in
Tasmanian and most of southern Australian waters. Off WA the species is reported from the western
Great Australian Bight off Eucla (128°E) and to the north off Shark Bay.

REFERENCE SPECIMENS. — AMS I.15970-009 (1 spec.); K71-06-04. AMS 1.15973-009 (1
spec.); K71-07-03. AMS 1.18838 -036 (6 spec.); K75-05-03. AMS I.18839-008 (6 spec.), AMS
1.18839-023 (4 spec.); K75-05-04. AMS I.19085-001 (1 spec.); K75-07-03. AMS 1.20099-004 (1
spec.); K77-23-12. AMS 1.201 18-032 (2 spec.); K77-13-10. AMS 1.20459-029 (2 spec.); K78-17-10.
AMS 1.28189-003 (1 spec.); K87-24-03. AMS I.29812-003 (1 spec.); K89-15-02.

REFERENCES. — Last et al. (1983); McMillan in Gomon et al. (1994); Iwamoto and Merrett
(1997); Iwamoto and Williams (1999).

Genus Mataeocephalus

DISTINGUISHING FEATURES. — Branchiostegal rays 6 or 7. Snout relatively long, somewhat flat-
tened, tipped with two tubercular scales; a series of coarse, modified scales along, and a naked groove
dorsally on each side behind, the leading edge. Mouth small, inferior, upper jaw length less than
one-third of HL. Outer gill rakers on first arch 0—S. Spinous ray of first dorsal fin slightly prolonged,
serrated along leading edge. Teeth on premaxillary in broad, short, truncated to slightly tapered band.
Periproct far removed from origin of anal fin. Scale spinules short, needlelike to lanceolate, in numer-
ous, more-or-less parallel rows.

REMARKS. — Two species in NSW,, including an undescribed species that is problematically
placed in Mataeocephalus. The genus is in need of revision and the characters in the generic diagnosis
apply only to the two NSW species. Species of the genus are usually caught in more tropical waters. It
is likely that central NSW is at the southern end of their normal distribution, as the two NSW species
(six specimens in total) were caught north of Newcastle. The small size of the two species also makes
capture by commercial-sized trawls difficult.

REFERENCES. — Iwamoto (1990); Iwamoto and Merrett (1997).

KEY TO THE SPECIES OF MATAEOCEPHALUS FROM NEW SOUTH WALES

la. Underside of snout mostly scaled; pelvic fin rays 7 (rarely 8); teeth in broadly tapered bands extending at least to
middlejofsyjawszibranchiostevaliraysiG: ayo etl: ces secs) Eke) seep ores oi cul t-te ten ene Mataeocephalus sp.

1b. Underside of snout almost entirely naked; pelvic fin rays 8—9; teeth in broad, short bands confined to front of mouth;
branchiostepalitayspiitas.. ate font oareeeucar) Chee nel eulsuinoe es lee ene Shay ky Re a M. acipenserinus

484 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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f

FIGURE 98. Mataeocephalus acipenserinus (Gilbert and Cramer, 1897). AMS 1.29753-009. From Kapala stn K89-17-06,
east of Crowdy Head, NSW, in 878-933 m.

Mataeocephalus acipenserinus (Gilbert and Cramer, 1897)
Fig. 98

DISTINGUISHING FEATURES. — ID II,8—10, spinous ray of 1D serrated along leading edge; P
119-125; V 8-9; total inner GR-I 6-8; scales below 2D 8-9, lat.1. about 40-42; pyloric caeca 13-19;
branchiostegal rays 7. Snout long, prominently protruding, length 39-44% HL; orbit diameter
29-34%; interorbital width 20-23%; mouth small, inferior, upper jaw length 20-28%; height 1D
50-63%. Underside of head naked except along front edge of snout; body scales with 4-6 parallel
rows of short conical spinules. Periproct oval to pear-shaped, situated midway between pelvic fin and
anal fin; small fossa of light organ anterior to anus. Overall color swarthy to pale brownish, darker
(bluish to violet) over abdomen; fins blackish to dusky.

SIZE. — To about 25 cm.

DISTRIBUTION. — Widespread in Pacific and Indian oceans, including Australia(NSW, WA), in
depths of about 600—900 m.

NSW CAPTURES. — Captured twice (five specimens) near Crowdy Head at somewhat greater
depths (823—933 m) than previously reported for the species.

REMARKS. — See Sazonov and Iwamoto (1992) for a detailed description and discussion of re-
lated taxa.

REFERENCE SPECIMENS. — AMS 1.29753-009 (4 spec.); K89-17-06. AMS 1.30304-007 (1
spec.); K89-17-09.

REFERENCES. — Sazonov and Iwamoto (1992); Iwamoto and Merrett (1997).

Mataeocephalus sp.
Fig. 99

DISTINGUISHING FEATURES. — ID II,8—10; 117-120; V 7 (rarely 6); total inner GR-I 7—8; scales
below 2D 5.5—7.5, lat.1. 31-35; pyloric caeca 16—18; branchiostegal rays 6. Snout of moderate length,
protruding, 32-36% HL; orbit diameter 24-29%; interorbital width 21-24%; mouth small, inferior,
upper jaw length 26-29%; height 1D 94-114%. Underside of head mostly scaled except for naked
median swath under snout; body scales covered with dense rows of lanceolate spinules. Periproct
small, anus closer to pelvic fin insertions than to anal fin origin. Few or no denticulations on spinous
ray of first dorsal fin.

SIZE. — To at least 21 cm.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 485

FIGURE 99. Mataeocephalus sp. AMS 1.29804-002. From Kapala stn K89-09-01, off Newcastle, NSW, in 896-960 m.

DISTRIBUTION. — Southwestern Pacific off New Caledonia and adjacent waters, and Australia
(Qld, NSW, WA), in depths of about 400 to almost 1000 m.

NSW CAPTURES. — Captured only once by Kapala, off Newcastle in 896-960 m.

REMARKS. — The six branchiostegal rays in this species set it apart from all other members of
this tribe (Malacocephalini). This species probably should be included in the genus Hyomacrurus
Gilbert and Hubbs, 1920. It awaits description by our Russian colleagues, Y. I. Sazonov and Y. N.
Shcherbachev.

REFERENCE SPECIMEN: AMS I.29804-002 (1 spec.); K89-09-01.

REFERENCES. — Iwamoto and Merrett (1997); Iwamoto and Williams (1999)

Genus Mesobius

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Anus situated in a broad oval periproct
area immediately anterior to anal fin. Body relatively deep, laterally compressed. Head smoothly
rounded, much deeper than wide, entirely covered with elongated scales with spinules aligned in 1-3
discrete longitudinal rows giving striated appearance to surface. Snout broadly rounded in lateral
view; no stout or sharp ridges on head. Chin barbel absent. Overall color of body and fins black, tail
somewhat paler.

REMARKS. — Two species, each sometimes taken in bathypelagic waters; only one species
known from NSW.

REFERENCES. — Hubbs and Iwamoto (1977); Arai (1979); Shcherbachev et al. (1979).

Mesobius antipodum Hubbs and Iwamoto, 1977
Fig. 100

DISTINGUISHING FEATURES. — 1D II,9—10; P 113-114; V 6—7; total GR-I (outer/inner) 7—10/13,
GR-II 13—14/12—-16; scales below 2D 9.5—10.5. Snout length 30-31% HL; orbit diameter 26-30%;
interorbital width 33-34%; suborbital width 14-16%; postorbital length 48-49%; upper jaw length
44-47%. Posttemporal region extending posterior to vertical through origin of pectoral fin base.

SIZE. — To 67 cm.

DISTRIBUTION. — Southern hemisphere, from South Atlantic off South Africa, through southern
part of Indian Ocean, southern coast of Australia (NSW, Vic., Tas., WA), to New Zealand, in
700-1300 m.

NSW CAPTURES. — Caught by Kapala on all mid-slope grounds south of Crowdy Head in 720—
1200 m. Relatively common. Mesobius antipodum was recorded from 168 Kapala stations (including
69% of all trawls deeper than 800 m). The average catch was about eight specimens, although many
trawls yielded more than 25 specimens.

486 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGurE 100. Mesobius antipodum Hubbs and Iwamoto, 1977. AMS 1.25095-007. From Kapala stn K84-20-05, off Broken
Bay, NSW, in 1170-1207 m.

REMARKS. — Shcherbachev et al. (1979) recorded Mesobius berryi Hubbs and Iwamoto, 1977,
the only congener of M. antipodum, from the Indian Ocean but not from off Australia. Those authors
provided a good comparison of the two species. Recently, Iwamoto and Williams (1999) reported one
specimen of M. berryi collected west of Cape Freycinet, Western Australia, in 1225-1240 m.

REFERENCE SPECIMENS. — AMS 1.20068-028 (1 spec.); K77-23-13. AMS 1.20098-004 (1
spec.); K77-23-07. AMS I.20485-003 (3 spec.); K77-23-06. AMS 25095-007 (1 spec.); K84-20-05.
AMS 1I.25290-011 (4 spec.); K84-20-03. AMS 1.25933-002 (1 spec.); K79-20-06.

REFERENCES. — McMillan in Gomon et al. (1994); Iwamoto and Williams (1999).

Genus Nezumia

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Snout pointed, slightly to extensively
protruding, tipped with paired, sometimes joined, thick tubercular scales. Chin barbel well developed.
Spinous ray of first dorsal fin serrated along leading edge. Anus closer to pelvic fin insertions than to
anal fin origin; periproct teardrop-shaped, a narrow connection to small dermal window of light organ
situated between pelvic fins. Underside of snout usually with naked area; suborbital shelf formed of
two rows of stout, coarsely modified scales. Teeth small, in band in both jaws, teeth in premaxillary
not reaching beyond posterior edge of maxillary process. Pyloric caeca usually not branched with
fewer than about 30 distal tips.

REMARKS. — More than 40 species known, but only four were found in NSW waters. In contrast,
seven were recorded from Western Australia by Iwamoto and Williams (1999). Several species previ-
ously classified under Nezumia (e.g., Iwamoto 1990) have been subsequently removed to other gen-
era, especially Kumba and Kuronezumia.

REFERENCES. — Iwamoto (1990); Iwamoto and Merrett (1997); Iwamoto and Williams (1999).

KEY TO THE SPECIES OF NEZUMIA FROM NEW SOUTH WALES

la. Pelvic fin rays 13-17; first dorsal fin pale or lightly dusky with a prominent black tip ........... N. propinqua
1b. Pelvic fin rays 9-12; first dorsal fin dusky to black, without a prominent black tip.................... 2

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 487

2a. Dermal window of light organ about on line with pelvic
fin insertions (Fig. 101a); dorsum rather uniformly pig-
mented, although area above abdomen often slightly
darker Be ena! o Ne eahale wh io] Loutsin's: Me, \eRie'g ee iw, ime N. coheni

2b. Dermal window of light organ well posterior to line con-
necting pelvic fin insertions (Fig. 101b); dorsal aspects
of trunk darker than comparable areas on tail, forming
complete dark band around trunk: ------- °°: 3

ADW

3a. Pelvic fin rays 9-10; sensory pores on underside of head
prominent (96-9 e - os es ahs cite es N. namatahi

3b. Pelvic fin rays 11-12; sensory pores on underside of
head small tet gel “see eRae ee ailiclrs, 16 Wet e> feute: Va ire N. kapala

Nezumia coheni lwamoto and Merrett, 1997
Fig. 102
FiGureE 101. Ventral view of belly of Nezumia spp. showing

DISTINGUISHING FEATURES. — 1D II,9—10; position of anterior dermal window (ADW) of light organ in
iPP=ig2seV 11 (carely 10 or’ 12); ‘total GR-1. © cohen! and) W. namaiahi
(outer/inner) 6—-9/9-11, GR-II 8—10/10-11;
scales below 1D 7-10, below 2D 7.0-8.5, lat.1. 34-40. Snout moderately protruding, length 30-34%
HL; orbit diameter 29-34%; interorbital width 20-26%; distance orbit to angle of preopercle
38-44%; postorbital length 39-45%; upper jaw length 30-34%; height 1 D about 80-95%. Underside
of head mostly scaled except for naked median swath under snout; body scales covered with dense
rows of lanceolate spinules. Periproct small, anus closer to pelvic fin insertion than to anal fin origin;
ADW about on line connecting insertions of pelvic fins. Dark band encircling trunk faint or lacking.

SIZE. — To more than 40 cm.

DISTRIBUTION. — Australia (NSW, Vic., SA), New Caledonia, and the Kermadec Is., in
710-1032 m.

NSW CAPTURES. — Recorded from 60 Kapala stations on all mid-slope grounds south of
Crowdy Head in 850-1200 m. Nezumia coheni was rare in catches south of Sydney, but two speci-
mens taken at one station in 1050 m off Gabo Island (37°40’S). Between Crowdy Head and Sydney,
the species was present in more than 40% of all trawls deeper than 900 m, with up to nine specimens
per trawl.

FIGURE 102. Nezumia coheni Iwamoto and Merrett, 1997. AMS 1.24181-003. From Kapala stn K83- 13-02, east of Newcas-
tle, NSW, in 960-988 m.

488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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REMARKS. — On first inspection, NV. coheni can easily be confused with the closely similar N.
kapala and N. namatahi, but those species have a prominent dark band completely encircling the
trunk. Nezumia namatahi has 9 or 10 pelvic fin rays, in contrast to the predominantly 11 of N. coheni,
and the anterior dermal window is well posterior to a line connecting the insertions of the pelvic fins.
Nezumia kapala has a somewhat shorter orbit to angle of preopercle distance (34-39% HL) than does
N. coheni, and its anterior dermal window is about on or (usually) well posterior to a line connecting
the insertions of the pelvic fins. All three species were captured together in a number of Kapala trawls;
of the 60 stations with N. coheni, 44 also contained N. kapala and/or N. namatahi.

REFERENCE SPECIMENS. — AMS I.21724-026 (2 paratypes); K79-20-15. AMS 1.24057-007 (2
spec.) and AMS 1.24057-008 (2 spec.); K83-09-04. AMS I.24173-006 (1 paratype); K83-14-06. AMS
1.24181-003 (1 spec.); K83-13-02. AMS 1I.24355-005 (1 spec.); K83-18-02. AMS 1I.24357-002 (2
spec.); K83-18-01. AMS 1.24993-007 (4 paratypes); K84-16-14. AMS 1.25127-002 (1 paratype);
K84-18-03. AMS 1.25264-003 (3 paratypes); K84-19-04. AMS I.25266-000 (2 spec.); K84-22-02.
AMS 1I.25290-006 (1 spec.); K84-20-03. AMS I.26247-005 (6 spec.); K85-21-04. AMS I.29340-007
(4 paratypes); K89-13-02. AMS 1.29741-004 (1 paratype); K87-25-06. AMS 1.29754-002 (1
paratype); K89-17-03. AMS 1.29761-004 (1 spec.); K89-19-02. AMS 1[.29823-013 (2 paratypes);
K89-13-01. AMS I.29827-003 (4 spec.); K89-15-01.

REFERENCES. — Iwamoto and Merrett (1997); Iwamoto and Williams (1999).

Nezumia kapala lwamoto and Williams, 1999
Fig. 103

DISTINGUISHING FEATURES. — 1D II,8-11; 118-122; V 11-12; total GR-I (outer/inner)
6-9/8—10, GR-IT 8—9/8—11; scales below 1D 8-12, below 2D 7.5—9.5, lat.1. 33-38. Snout moderately
protruding, length 27-34% HL; orbit diameter 29-33%; interorbital width 18-25%; distance orbit to
angle of preopercle 34-39%; postorbital length 40-45%; upper jaw length 26-31%; height 1D about
86—-109%. Underside of head mostly scaled except for naked median swath under snout; body scales
covered with dense rows of lanceolate spinules. Periproct small, anus closer to pelvic fin insertions
than to anal fin origin; anterior dermal window about on, or usually well behind, line connecting inser-
tions of pelvic fins. A prominent dark band encircling trunk.

SIZE. — To about 41 cm.

FIGURE 103. Nezumia kapala |wamoto and Williams, 1999. AMS 1.24178-003. From Kapala stn K83-14-08, off Broken
Bay, NSW, in 978-1006 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 489

DISTRIBUTION. — Australia (NSW, Tas., WA) in 842-1243 m. Can be expected off Vic. and
SA.

NSW CAPTURES. — Nezumia kapala was not distinguished from N. namatahi in Kapala field re-
cords. Based on 15 AMS registrations, N. kapala was caught by Kapala between 32°01'S (Crowdy
Head) and 35°30'S (se. of Ulladulla) in 930-1100 m. Either or both species were caught in 135
Kapala stations between Crowdy Head and Gabo Island; most catches were between 4 and 8 speci-
mens.

REMARKS. — Nezumia kapala and N. namatahi are very similar in overall appearance, and as
they are often caught together, they are susceptible to being confused with one another. The pelvic ray
counts, however, readily distinguish the species (11 or 12 in N. kapala, 9 or 10 in N. namatahi).
Nezumia namatahi also has more broadly lanceolate to shield-shaped scale spinules and larger sen-
sory pores under the head. Specimens of N. namatahi and N. kapala in AMS were collected without
knowledge of their distinction. Subsequently, both species were found together in five of the stations
represented in the collection. This suggests that there is a high probability that both species, recorded
as “N. namatahi,” were present in a significant proportion of the 135 Kapala stations

REFERENCE SPECIMENS. — AMS 21724-005 (1 paratype); K79-20-15. AMS 1.24057-001 (4
paratypes); K83-09-04. AMS 1I.24059-006 (4 paratypes), AMS 1I.24059-021 (2 spec.); K83-09-02.
AMS 1.24060-023 (1 paratype); K83-09-01. CAS 200228, formerly AMS 1.24150-006 (1 paratype);
K83-13-01. AMS 1.24173-009 (1 spec.); AMS I.24173-010 (1 spec.); AMS 1.24173-012 (1 spec.);
K83-14-06. AMS 1.24178-003 (1 spec.); K83-14-08. AMS _ 1.24993-008 (holotype), AMS
1.24993-010 (1 paratype); K84-16-14. AMS I.25127-006 (1 spec.); K84-18-03. AMS I.25264-002 (4
paratypes); K84-19-04. AMS 1.25266-008 (2 paratypes); K84-22-02. AMS I.29741-005 (1 spec.);
K87-25-06. AMS 1.29754-007 (1 spec.); K89-17-03. AMS 1.29761 -002 (1 spec.); K89-19-02. AMS
1.29797-006 (2 spec.); K89-12-05. AMS 1.29827-004 (3 spec.); K89-15-01.

REFERENCES. — McMillan in Gomon et al. 1994; “darknose whiptail,” in part; Iwamoto and
Williams (1999).

Nezumia namatahi McCann and McKnight, 1980
Fig. 104

DISTINGUISHING FEATURES. — 1D II,9—10, rarely 11; 118-121; V 9-10, usually 10; total GR-I
(outer/inner) 5—8/7—9, GR-IT 7—9/8—10; scales below 1D 8—11, usually 9-10, below 2D 7.5—9.5, lat.1.
32-38; pyloric caeca about 21—22. Snout length 29-33% HL; orbit diameter 32-38%; interorbital
width 19-24%; distance orbit to angle of preopercle 33-37%; postorbital length 37-43%; upper jaw
length 24-30%; barbel length 14-30%; height 1D about 90—121%. Underside of snout broadly naked,
sensory pores on head prominent; body scales covered with broadly lanceolate to shield-shaped spi-
nules in parallel to slightly convergent rows. Periproct small, anus closer to pelvic fin insertions than
to anal fin origin; ADW usually well behind line connecting insertions of pelvic fins. A prominent
dark band encircling trunk.

SIZE. — To at least 36 cm.

DISTRIBUTION. — Australia (NSW, Tas., Vic.?, to 138°E in SA) and New Zealand, in
700-1170 m.

NSW CAPTURES. — Based on 10 AMS registrations, the species was caught between 32°28'S
(ne. of Port Stephens) and 34°55'S (Jervis Bay) in 730-1150 m. (See NSW Captures section for N.
kapala for discussion of both species.)

REMARKS. — Nezumia toi McCann and McKnight, 1980 is a synonym of this species and was
based on an immature specimen in rather poor condition. (See Remarks section for N. kapala for dis-
cussion of both species.)

490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 104. Nezumia namatahi McCann and McKnight, 1980.

REFERENCE SPECIMENS. — AMS 21724-002 (3 spec.); K79-20-15. AMS I.23460-001 (1 spec.);
stn NZOI U-222; ne. of Newcastle, 32°49’S, 152°49'E; 1040-1075 m; 9 Oct. 1982. AMS I.24057-009
(1 spec.); K83-09-04. AMS 1I.24059-010 (1 spec.), AMS 1.24059-022 (1 spec.); K83-09-02. AMS
1.24060-017 (1 spec.); K83-09-01. AMS 1.24980-004 (3 spec.); K84-16-15. AMS I. 25290-010 (1
spec.); K84-20-03. AMS 1.29340—006 (2 spec.); K89-13-02. AMS 1.29754-005 (1. spec.);
K89-17-03. AMS I.29823-014 (1 spec.); K89-13-01.

REFERENCES. — McCann and McKnight (1980); Iwamoto and Williams (1999).

Nezumia propinqua (Gilbert and Cramer, 1897)
Fig. 105

DISTINGUISHING FEATURES. — 1D II,10—12; 119-122; V 13-17; total GR-I (outer/inner)
8—10/8—10, GR-II 7—9/9-10; scales below 1D 10-13, below 2D 8.5—10, lat.1. 36-42; pyloric caeca
about 21—28. Snout length 29-34% HL; orbit diameter 30-34%; interorbital width 20-25%; distance
orbit to angle of preopercle 30-39%; postorbital length 41-45%; upper jaw length 31-35%; barbel
length 15-23%; height 1D about 97—1 13%. Underside of snout, suborbital, and lower jaw naked, sen-
sory pores on naked areas small but prominent; body scales covered with spinules in 10—12 parallel to
slightly convergent rows. Periproct large, situated about midway between pelvic fin insertions and
anal fin origin; ADW slightly in advance of line connecting insertions of pelvic fins. First dorsal fin
with prominent black tip; no dark band encircling trunk. (After Iwamoto and Williams 1999.)

SIZE. — To about 25 cm.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 491

FiGuRE 105. Nezumia propinqua (Gilbert and Cramer, 1897). AMS 1.27722-002. From Kapala stn K88-08-09, east of New-
castle, NSW, in 704-750 m.

DISTRIBUTION. — Widespread in tropical waters of Pacific and Indian oceans, including Austra-
lia (Qld, NSW, WA). Capture depths about 400—1100 m.

NSW CAPTURES. — Recorded from 43 mid-slope stations (85 specimens) between Crowdy
Head and Batemans Bay in 660—1 100 m. Most captures were north of Sydney in 800—900 m; 38 of the
43 stations were with small-meshed nets, suggesting that the small N. propinqua was seldom retained
in the larger-meshed trawls.

REMARKS. — There is some uncertainty as to the status of this and two closely similar species, N.
condylura (Jordan and Gilbert, 1904) and N. evides (Gilbert and Hubbs, 1920). Iwamoto and Williams
(1999) and Sazonov and Iwamoto (1992) briefly discuss the problems.

REFERENCE SPECIMENS. — AMS 1I.19859-009 (3 spec.); K76-24-04. AMS I.19860-018 (8
spec.); K76-24-03. AMS I.20484-003 (1 spec.); K77-21-01. AMS 1.20485-005 (1 spec.); K77-23-06.
AMS 1I.23710-002 (1 spec.); K79-20-04. AMS I.24037-005 (1 spec.); K78-26-16. AMS 1.24613-003
(2 spec.); K75-05-05. AMS 1.24979-001 (2 spec.); K84-16-04. AMS 1.24980-002 (5 spec.);
K84-16-15. AMS I.24981-001 (1 spec.); K84-17-04. AMS 1.24989-002 (1 spec.); K84-17-01. AMS
1.24990-002 (1 spec.); K84-16-13. AMS 1.26000-003 (1 spec.); K86-01-08. AMS 1I.26002-003 (1
spec.); K86-01-06. AMS I.26245-014 (1 spec.); K86-01-07. AMS I.27722-002 (1 spec.); K88-08-09.
AMS 1.29591-002 (1 spec.); K88-12-03. AMS 1.29601-002 (1 spec.); K88-08-08. AMS 1.29743-001
(2 spec.); K89-20-01. AMS 1.29749-001 (1 spec.); K89-17-02. AMS 1.29753-004 (1 spec.);
K89-17-06. AMS I.29754-006 (1 spec.); K89-17-03. AMS 1.29757-001 (3 spec.); K89-15-03. AMS
1.29762-006 (1 spec.); K89-12-03. AMS 1I.29797-007 (2 spec.); K89-12-05. AMS 1.29803-002 (1
spec.); K89-07-01. AMS I.29805-002 (1 spec.); K87-24-04. AMS 1.2981 1-004 (1 spec.); K89-08-01.
AMS 1.29823-004 (5 spec.); K89-13-01. AMS 1.30304-002 (1 spec.); K89-17-09.

REFERENCE. — Sazonov and Iwamoto (1992); Iwamoto and Merrett (1997); Iwamoto and Wil-
liams (1999).

Genus Odontomacrurus

DISTINGUISHING FEATURES. — Branchiostegal rays 6. Long spinous ray of first dorsal fin
smooth along leading edge. Mouth large, jaws armed with fanglike teeth in one row. Chin barbel ab-
sent. Anus midway between pelvic fin insertions and anal fin origin, preceded by a small fossa of light
organ. Large, prominent sensory pores on head. Lateral line in two parts, anterior section short,
dorsolateral, ending below hind margin of first dorsal fin.Swim bladder very small. Color overall
black to swarthy.

492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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REMARKS. — One widespread bathypelagic species. Known only from tropical and subtropical
waters.
REFERENCES. — Norman (1939); Marshall (1964).

Odontomacrurus murrayi Norman, 1939
Fig. 106

DISTINGUISHING FEATURES. — As for genus.

SIZE. — To about 55 cm.

DISTRIBUTION. — Widely distributed in Atlantic and Indian oceans; in western Pacific previ-
ously recorded only from South China Sea, but now known from off southeastern Australia (NSW,
Tas.).

NSW CAPTURES. — Specimens were caught by Kapala in each of four midwater trawl stations
between Port Stephens and Wollongong. Capture depths were 0—900 m in oceanic waters deeper than
2000 m.

REMARKS. — This species and Cynomacrurus piriei are among the few strictly bathypelagic
species of grenadier. Cynomacrurus differs in having the anus immediately before the anal fin and a
posterior midlateral section of the lateral line, and in lacking an abdominal fossa of the light organ.
Odontomacrurus murrayi is a broadly distributed species but rarely captured, probably because large
midwater trawls are seldom towed in oceanic waters (the Kapala specimens were taken when trawling
for bathypelagic and mesopelagic fishes for AMS).

REFERENCE SPECIMENS. — AMS I.20064-023 (1 spec.); K77-18-01. AMS 1.20314-047 (1
spec.); K77-24-10. AMS 1.203 15-050 (1 spec.); K77-24-11. AMS I.21369-004 (1 spec.); K79-19-07.

REFERENCES. — Norman (1939); Marshall (1964); Iwamoto (1970).

Genus Sphagemacrurus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Snout blunt, upturned, tip and lateral an-
gles armed with spiny tubercular scales. Mouth cleft moderately to steeply oblique. Chin barbel pres-
ent. Spinous ray of first dorsal fin serrated along leading edge. Ventral aspects of body shifted forward
so that pelvic fin origin usually under opercle, anal fin origin under first dorsal fin, gill membranes
united under preopercle. Broad, naked periproct region abutting anal fin origin; small fossa of light or-
gan between pelvic fins but well posterior to pelvic fin base. Underside of snout variously naked; sub-

FIGURE 106. Odontomacrurus murrayi Norman, 1939. AMS 1.20064-023. From Kapala stn K77-18-01, midwater trawl in
0-900 m, 42 n. mi. ese. of Sydney, NSW.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 493

orbital shelf formed by two rows of stout, coarsely modified scales. Body scales covered with several
rows of short, slender, conical spinules in parallel to slightly divergent rows. Teeth small, in narrow to
moderately wide band in both jaws, teeth in premaxillary not reaching beyond posterior edge of
maxillary process. Pyloric caeca usually less than 30.

REMARKS. — Six species recognized, but only one presently known from NSW.
Sphagemacrurus pumiliceps (Alcock, 1894) could be expected, as the species is known from Qld,
WA, New Caledonia, and other areas in the Indian and Pacific oceans.

REFERENCES. — Weber and de Beaufort (1929); Iwamoto (1990); Iwamoto and Williams
(1999).

Sphagemacrurus richardi (Weber, 1913)
Fig. 107

DISTINGUISHING FEATURES. — 1D _ II,9-11; 117-120; V 8-11; total GR-I (outer/inner)
9—-12/11-13, GR-II 9-11/11—13; scales below 1D 11-14, below 2D 8.5-9.5, lat.1. 37-43; pyloric
caeca about 10. Snout length 30-37% HL; internasal width 28-33%; interorbital width 28-32%; orbit
diameter 31—36%; distance orbit to angle of preopercle 39-44%; postorbital length 36-41%; upper
jaw length 35-40%; barbel length 13-16%; height 1D about 85—100%; distance outer pelvic ray to
anal fin origin about 30-40%. Body scales covered with short, fine spinules in 7—9 parallel rows.
Periproct large, immediately before anal fin origin and spanning about half distance to pelvic fin in-
sertion; ADW extending forward from periproct. Head relatively pale along sides, eye ring promi-
nent; trunk and tail darker; abdomen and chest dark with violet hue; first dorsal, pectoral, and anal fins
dusky, pelvic fin blackish.

SIZE. — To about 23 cm.

DISTRIBUTION. — Indonesia and Australia (NSW).

NSW CAPTURES. — Uncommon in Kapala catches; caught on 18 occasions (56 specimens)
north of Sydney at 880-1100 m depth. A species of small adult size, most were caught in trawls with
small-meshed codends and were probably more abundant than their capture rate suggests.

REMARKS. — The species appears to be closely similar to S. decimalis (Gilbert and Hubbs, 1920)
from the Philippines, but that species has a lower gill raker count (about 7 on lower limb of first arch).
Sphagemacrurus pumiliceps (Alcock, 1894) has somewhat higher pelvic fin ray counts (11—14), a
longer barbel (16-22% HL), and narrower interorbital (23—26% HL).

REFERENCE SPECIMENS. — AMS I.26000-002 (1 spec.); K86-01-08. AMS 1.26247-001 (1
spec.); K85-21-04. AMS I.27720-001 (1 spec.); K88-08-04. AMS I.27721-001 (3 spec.); K88-08-07.
AMS I.28900-001 (3 spec.); K88-17-06. AMS 1.28988-002 (5 spec.); K89-09-03. AMS 1.29298-001

FiGuRE 107. Sphagemacrurus richardi (Weber, 1913). AMS 1.27721-001. From Kapala stn K88-08-07, east of Newcastle,
NSW, in 1006-1079 m.

494 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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(4 spec.); K88-20-03. AMS 1.29605-006 (5 spec.); K89-09-07. AMS 1.29750-002 (6 spec.);
K89-17-04. AMS I.29753-01 1 (2 spec.); K89-17-06. AMS 1.29799-002 (6 spec.) and I.29799-007 (7
spec.); K89-06-02. AMS I.29809-001 (6 spec.); K88-20-01. AMS 1.29823-012 (1 spec.); K89-13-01.
AMS I.29827-005 (8 spec.); K89-15-01.

REFERENCES. — Weber (1913); Weber and de Beaufort (1929); Iwamoto (1990).

Genus Trachonurus

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Snout rounded, lacking a prominently
protruding terminal tubercle. Chin barbel present. Broad, naked black periproct region extending
most of (relatively short) distance between pelvic and anal fins. Spinous ray of first dorsal fin flexible,
smooth along leading edge. Origin of pelvic fin usually behind pectoral fin base. Head almost fully
scaled except for patches or single scales on branchiostegal and gular membranes in some species;
suborbital vertical in most, with smoothly rounded contours, lacking sharp or coarsely scaled ridges.
Body scales covered with short, conical spinules in somewhat quincunx pattern. Teeth in narrow band
in upper jaw with outer series usually somewhat enlarged, teeth in lower jaw in 2 or 3 rows or narrow
band. Pyloric caeca usually less than 15. Color overall black or brown or gray.

REMARKS. — At least six species, two of which are recorded from NSW. Iwamoto and Williams
(1999) were uncertain as to the identification of one specimen (AMS I.27718-010) from southern
NSW off Ulladulla. Its characters did not agree
well with the characters of 7. sentipellis and may
represent another species. The key provided be-
low is adapted from Iwamoto and Williams
(1999:212), but does not include 7. yiwardaus
Iwamoto and Williams, 1999, a species so far
known only from WA and SA.

REFERENCES. — Iwamoto and McMillan
(1997); Iwamoto and Merrett (1997); Iwamoto
and Williams (1999).

KEY TO THE SPECIES OF TRACHONURUS FROM
NEW SOUTH WALES

la. Grooved lateral line present; body scales large, 8 or 9
rows between pelvic fin base and gill cover (Fig. 108a)
SE se ee ariiee rab anece rt iol ichlo/ co. Sie tome c T. sentipellis

1b. Grooved lateral line absent; body scales relatively small,
10—14 rows between pelvic fin base and gill cover (Fig.
108b)2 eee ee T. gagates

Trachonurus gagates wamoto and McMillan,
1997
Fig. 109

DISTINGUISHING | FEATURES. — Grooved
lateral line absent. Chin barbel short, length 4-8%
of HL. Scale rows between pelvic fin and gill

cover 10-14. Color uniformly black to dark

brown. FiGURE 108. Diagrammatic lateral views of Trachonurus
spp. showing method of counting scale rows between pelvic

SIZE. — To about 48 cm. fin base and gill cover in (a) T. sentipellis and (b) T. gagates.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 495

Ficure 109. Trachonurus gagates Iwamoto, McMillan, 1997. AMS I.24059-009. From Kapala stn K83-09-02, east of Bro-
ken Bay, NSW, in 933-969 m.

DISTRIBUTION. — Australia (Qld, NSW, Vic., Tas., SA, WA) and New Zealand, in 435—1200 m.

NSW CAPTURES. — Recorded in 65 Kapala stations from Crowdy Head to se. of Gabo Island;
depth range 890-1200 m. An average of two specimens was caught in the 65 stations; most were
caught deeper than 1000 m where 47% of trawls contained 7. gagates.

REMARKS. — This relatively large, dark species with small body scales is readily distinguished
from its congeners by the absence of a grooved lateral line. One specimen (AMS I.20307-067) was
taken by midwater trawl in oceanic waters.

REFERENCE SPECIMENS. — AMS 1.20307-067 (1 spec.); K77-24-03). AMS _ 1.24059-009
(holotype); K83-09-02. AMS 1.24157-002 (1 spec.); K83-12-04. AMS 1.24173-007 (3 spec.) and
AMS 1.24173-011 (7 spec.); K83-14-06. AMS I.24178-004 (1 spec.); K83-14-08. AMS I.24355-001
(1 spec.); K83-18-02. AMS 1.24356-005 (3 spec.) and AMS I.24356-007 (1 spec.); K83-14-05. AMS
1.24357-001 (1 spec.); K83-18-01. AMS 1.24451-002 (2 spec.); K83-14-09. AMS 1.24625-003 (3
spec.); K84-06-06.

REFERENCES. — Paxton et al. (1989)(as T. villosus); Iwamoto and McMillan (1997).

Trachonurus sentipellis Gilbert and Cramer, 1897
Fig. 110

DISTINGUISHING FEATURES. — Grooved lateral line present. Body scales relatively large,
coarsely covered with stout, erect spinules, 26—34 lat.|. scales over distance equal to predorsal length,
4-7 scale rows below midbase of first dorsal fin, 5—7 below origin of second dorsal; 8 or 9 between

FIGURE 110. Trachonurus sentipellis (Gilbert and Cramer, 1897). AMS I.28100-003. From Kapala stn K88-08-06, east of
Tuncurry, NSW, in 1024-1079 m.

496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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pelvic fin base and gill cover. Small scale patch or none on gular membrane, few or no scales on
branchiostegal rays. Teeth in both jaws small, outer premaxillary series scarcely enlarged. Chin bar-
bel moderate, 9-14% of HL. Total GR-IT (outer) 10—14. Pyloric caeca short, thick, 9-13. Color uni-
formly black to dark brown.

SIZE. — To 31+ cm.

DISTRIBUTION. — Hawaiian Is., Australia (NSW, WA), and New Caledonia, in 500-1136 m.

NSW CAPTURES. — Three specimens taken by Kapala in 940-1130 m off Crowdy Head at the
northern end of the mid-slope grounds.

REMARKS. — Iwamoto and Williams (1999) listed four specimens as uncertain variants of this
species. The four showed differences in scale distribution and spinulation, gill-raker and scale-row
counts, and some proportional measurements, compared with their other specimens, indicating the
possibility of more than one species being involved. One of the uncertain variants was from NSW
(AMS I.27718-010, off Ulladulla in 1150 m).

REFERENCE SPECIMENS. — AMS 1.24462-003 (1 spec.); K83-15-02. AMS 1.28100-003 (1
spec.); K88-08-06. AMS I.29808-001 (1 spec.); K89-06-04.

REFERENCES. — Iwamoto and Merrett (1997); Iwamoto and Williams (1999).

Genus Ventrifossa

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Snout moderately pointed to bluntly
rounded, lacking thickened tubercular scales at lateral angles and (in most species) tip of snout. Chin
barbel well developed. Spinous ray of first dorsal fin finely serrated along leading edge in NSW spe-
cies. Anus closer to pelvic fin insertions than to anal fin origin; periproct teardrop-shaped, a narrow
connection to small dermal window of light organ situated between pelvic fins. Underside of snout
fully and uniformly scaled; suborbital ridge rounded, without coarsely modified scales. Teeth in
bands in both jaws, outer premaxillary teeth enlarged, with tooth band extending beyond posterior
edge of maxillary process. Pyloric caeca more than 30. Color often silvery along sides of head and
body; lips usually black; leading edge of snout, suborbital shelf, and dorsal snout ridges in most spe-
cies dark, or terminal snout scute blackish.

REMARKS. — More than 25 species. Only three species recorded from NSW waters, although
seven are known from Western Australia (Iwamoto and Williams 1999).

REFERENCES. — Iwamoto (1990); Iwamoto and Merrett (1997); Iwamoto and Williams (1999).

KEY TO THE SPECIES OF VENTRIFOSSA FROM NEW SOUTH WALES

la. A tubercular scale at tip of snout; suborbital shelf narrowly constricted anteriorly; lateral line scales over distance equal

to’ predorsalilengthtofjheadiG4=79 00... 25 ocsgs es es Gis aye aide ae ss ees | Ge V. johnboborum
1b. No tubercular scale at tip of snout; suborbital shelf not especially constricted anteriorly; lateral line scales over distance

equalitojpredorsalllengthrofiheadilessithan'Sik S75 5% 3.2 Se se ee ee ee 2 eS eee 2
2a. A prominent blotch or dark streak across first dorsal fin; pelvic fin rays 8 or 9, usually 8 . . . Ventrifossa nigrodorsalis
2b. First dorsal fin dark overall, without blotch or streak; pelvic fin rays 9 or 10, usually9 ............ V. paxtoni

Ventrifossa johnboborum Iwamoto, 1982
Fig. 111

DISTINGUISHING FEATURES. — 1D _ II,9-11; il7-i23; V 8-9; total GR-I (outer/inner)
9—12/13-15, GR-IT 12—15/12—14; scales below 1D about 12—16, below 2D about 9-12, lat.1. 64—75.
Snout moderately protruding, length 28-31% HL; orbit diameter 29-35%; interorbital width
25-30%; distance orbit to angle of preopercle 41-44%; postorbital length 41-44%; upper jaw length
36-41%; barbel length 7-13 (18)%; height 1D about 54-68%. Suborbital shelf extremely narrow an-

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 497

FiGurE 111. Ventrifossa johnboborum Iwamoto, 1982. AMS I.25126-001. From Kapala stn K84-18-06, east of Sydney,
NSW, in 914-924 m.

teriorly, but broadening posteriorly. Body scales small, densely covered with small needlelike spi-
nules. Spinous second ray of first dorsal fin with finely serrated leading edge. Tip of snout (and
sometimes leading edge) blackish or dark dusky, other head ridges not marked. Lining of mouth dark.

SIZE. — To about 48 cm.

DISTRIBUTION. — Western Indian Ocean, east to Australia (Qld, NSW, WA) and New Caledo-
nia, north to Philippines and South China Sea, and in the southeastern Pacific at Sala-y-Gomez Ridge.
Depth range about 400-1100 m.

NSW CAPTURES. — Recorded from 23 Kapala stations (88 specimens) between Crowdy Head
and Ulladulla, all but one station north of Sydney. Depth range 680—980 m, plus a single capture in
1070 m. Not commonly caught, V. johnbororum was present in only 20 of the 98 trawls in
700-1000 m north of Sydney. Most trawls caught less than five specimens, but 25 were taken in one
station off Port Stephens.

REMARKS. — The northerly NSW distribution conforms to the generally tropical occurrence of
the species. However, the NSW capture depths were somewhat greater than previously reported
(412-855 m). Some questions remain as to the taxonomic status of the various populations of this ap-
parently widespread species. Ventrifossa fusca Okamura, 1982 from the Kyushu-Palau Ridge and V.
misakia Jordan and Gilbert, 1904 from Japan are closely related to this species, and the three compose
the subgenus Sokodara.

REFERENCE SPECIMENS. — AMS I.19860-014 (1 spec.); K76-24-03. AMS 1.25126-001 (1
spec.); K84-18-06. AMS I.27647-003 (1 spec.); K87-16-02. AMS 1.28189-004 (1 spec.); K87-24-03.
AMS 1.29746-003 (2 spec.); K88-21-03. AMS I. 29747-004 (3 spec.); K88-21-02. AMS I.29749-007
(1 spec.); K89-17-02. AMS 1.29756-003 (3 spec.); K89-15-04. AMS 1.29762-005 (3 spec.);
K89-12-03. AMS I.29806-001 (1 spec.); K89-09-06. AMS I.29811-002 (1 spec.); K89-08-01. AMS
1.29813-007 (1 spec.); K89-06-05. AMS 1.30737-002 (4 spec.); K87-24-02.

REFERENCES. — Sazonov and Iwamoto (1992); Iwamoto and Merrett (1997); Iwamoto and Wil-
liams (1999).

Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920
Fig. 112

DISTINGUISHING FEATURES. — 1D II,9—11; 118-123; V 8-9 (usually 8); total GR-I outer/inner)
8—12/13—16, GR-II 13—15/12—14; scales below 1D about 7-10, below 2D 7.5—10, lat.1. 39-42. Snout
slightly protruding, length 26-33% HL; orbit diameter 29-35%; interorbital width 24-30%; distance

498 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 112. Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920. From Kapala stn K83-01-08, off Sydney, NSW, in
490-570 m.

orbit to angle of preopercle 38-45%; postorbital length 40-46%; upper jaw length 37-45%; barbel
length 16-27%; height 1D about 64-96%. Suborbital shelf uniformly broad. Body scales thin, rather
deciduous, covered with small conical spinules in quincunx pattern. Spinous second ray of first dorsal
fin with finely serrated leading edge. Dorsal surfaces of trunk and tail dark, contrasting with silvery
(when fresh) or pale (when preserved) ventral body surfaces. Prominent black blotch or streak across
anterior portion of first dorsal fin. Leading edge of snout, supranasal ridge (but not median nasal
ridge), and suborbital shelf blackish.

SIZE. — To about 30 cm.

DISTRIBUTION. — Widespread in the western Pacific, from the Philippines, South China Sea,
and Indonesia south to New Caledonia and Australia (Qld, NSW, WA). Depth range about
300-800 m.

NSW CAPTURES. — Relatively common in upper slope depths off central and northern NSW,
particularly in 500-700 m. Captured by Kapala in 71 trawls on all grounds between the Qld-NSW
border and Jervis Bay (35°00'S) in 360-790 m. Recently collected south of Bermagui to 36° 46'S. Be-
tween 10 and 20 specimens were caught at most Kapala stations with small-meshed nets. Because of
its small size, few were caught with large-meshed trawls.

REMARKS. — An apparently widespread, highly variable species deserving further study (see
Iwamoto and Williams 1999).

REFERENCE SPECIMENS. — AMS I.15987-014 (1 spec.); K71-11-09. AMS I.18839-013 (1
spec.); K75-05-04. AMS I.20301-025 (1 spec.); K77-13-12. AMS 1.20459-015 (17 spec.) and
1.20459-027 (4 spec.); K78-17-10. AMS 1.20518-008 (20 spec.) and 1.20518-018 (2 spec.);
K78-09-05. AMS I1.21669-004 (3 spec.); K77-16-16. AMS 1.21805-002 (25 spec.); K77-23-09. AMS
1.21806-002 (3 spec.); K77-07-10. AMS 1.23710-001 (4 spec.); K79-20-04. AMS 1.24852-009 (1
spec.); K84-15-03. AMS I.29535-005 (1 spec.); K79-15-01. CAS 214043 (6 spec.); off Bermagui
(36°46'S, 150°21’'E); 530-549 m; FV Shelley H, 15 Feb. 2000. CAS 214044 (6 spec.); off Bermagui
(36°12'S, 150°24’E); 421 m; FV Shelley H, | Mar. 2000.

REFERENCES. — Gilbert and Hubbs (1920); Iwamoto and Merrett (1997); Iwamoto and Wil-
liams (1999).

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 499

Ventrifossa paxtoni Twamoto and Williams, 1999
Fig. 113

DISTINGUISHING FEATURES. — 1D IT,9—11; 120-125; V 9-10 (usually 9); total GR-I outer/inner)
10—13/15—18, GR-II 14—18/15—18; scales below 1D about 8—12, below 2D 7.5—9.0, lat.l. 37-50.
Snout rather blunt, length 24-31% HL; orbit diameter 27-38%; interorbital width 24-30%); distance
orbit to angle of preopercle 42-48%; postorbital length 43-53%; upper jaw length 43-50%; barbel
length 24-38%; height 1D about 63-73%. Suborbital shelf uniformly broad. Body scales covered
with small conical spinules in subparallel rows. Spinous second ray of first dorsal fin with finely ser-
rated leading edge. Color overall swarthy, especially dark over head, chest and abdomen. Fins all
black or blackish. Leading edge of snout, lateral nasal ridges, and suborbital shelf black, median nasal
ridge dusky to blackish.

SIZE. — To about 43 cm.

DISTRIBUTION. — Australia (Qld, NSW, WA) and the New Caledonian region, in about
800-1100 m.

NSW CAPTURES. — Captured at 27 Kapala stations between Crowdy Head and Sydney, with ad-
ditional single captures off Nowra and Ulladulla; depth range 825—1050 m. Caught in about 20% of
all trawls in 800-1100 m north of Sydney. Overall average catch about four per trawl, with highest
catch numbers at stations north of Newcastle (up to 25 specimens).

REMARKS. — Ventrifossa paxtoni is a dark, blunt-snouted species similar to V. macropogon, a
widespread species from the western North Atlantic, Western Australia, and New Caledonia.
Ventrifossa paxtoni, however, has a somewhat shorter, thinner barbel, fainter median nasal streak, and
higher gill raker counts. Ventrifossa saikaiensis Okamura, 1984 is also closely similar but has some-
what lower counts of pelvic fin rays (8 or 9), slightly higher counts of outer gill rakers (12-15), and
lacks darkly marked head ridges, including the leading edge of the snout.

REFERENCE SPECIMENS. — AMS 1.20099-006 (2 spec.); K77-23-12. AMS 1.24150-004 (1
spec.); K83-13-01. AMS 1I.24990-001 (1 paratype); K84-16-13. AMS I.26981-005 (5 paratypes);
K87-02-01. AMS 1I.27647-002 (holotype) and AMS 1.27647-004 (1 paratype); K87-16-02. AMS
1.28189-005 (2 paratypes); K87-24-03. AMS 1.29753-008 (4 paratypes); K87-17-06. AMS
1.29797-005 (4 spec.); K89-12-05. AMS I.29805-001 (1 paratype); K87-24-04. AMS 1.30737-001 (4
spec.); K87-24-02.

REFERENCES. — Iwamoto and Merrett (1997); Iwamoto and Williams (1999).

FicurE 113. Ventrifossa paxtoni Iwamoto and Williams, 1999. AMS 1.24150-004. From Kapala stn K83-13-01, from off
Broken Bay, NSW, in 988-1015 m.

500 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SUBFAMILY TRACHYRINCINAE

DISTINGUISHING FEATURES. — Branchiostegal rays 7. Outer gill slit wide and free, not restricted
by folds of skin connecting upper and lower limbs of gill arch. Dorsal fins two, closely approximated,
the first short-based and armed with a flexible leading spinous ray; the second long and continuous to
end of tail. Anal fin long, height usually somewhat lower and length slightly shorter than second dor-
sal fin. A rudimentary caudal fin sometimes developed. Heavy scutelike scales forming long ridges
along dorsal and ventral margins lateral to median fins. No light organ.

REMARKS. — Two genera, /diolophorhynchus (monotypic) and Trachyrincus (with six species,
one of which is found off NSW). /diolophorhynchus has leathery head ridges and scales that have low,
flattened, or no spinules on exposed fields; a midlateral row of enlarged scutes on trunk; pelvic fin rays
3 or 4; no chin barbel; and no posttemporal pit. The single species, /. andriashevi Sazonov, 1981, may
occur off NSW at depths greater than trawled by the Kapala.

REFERENCES. — Iwamoto (1990); McMillan (1995).

Genus Trachyrincus

DISTINGUISHING FEATURES. — V 6 or 7. Snout strongly supported and pointed. A sensory pit in
temporal region of head. Small chin barbel present. Scales on head and body covered with stout spi-
nules. Pyloric caeca bifid.

REFERENCE. — McMillan (1995).

Trachyrincus longirostris (Gunther, 1878)
Fig. 114

DISTINGUISHING FEATURES. — Snout long, 41-46% of HL, ventral length 34-39%; upper jaw
length 25—28%; pyloric caeca 35—62. Color pale brownish to grayish.

SIZE. — To about 50 cm.

DISTRIBUTION. — Australia (NSW, Vic.), New Zealand, and southern Africa, in about
1100-1400 m.

NSW CAPTURES. — Caught at four Kapala stations (five specimens) off Port Stephens, Broken
Bay (2) and Gabo Island in 1050-1200 m. Possibly more abundant in depths greater than 1200 m.

REMARKS. — Trachyrincus aphyodes McMillan, 1995 from New Zealand waters is closely sim-
ilar to T. Jongirostris, but may be distinguished by its more numerous pyloric caeca (1 19-211), some-
what longer upper jaw (29-34% of HL), and shorter ventral snout length (26-34%). Trachyrincus
longirostris is a smaller species, attaining about 50 cm, compared with 96 cm in T. aphyodes.

REFERENCE SPECIMENS. — AMS 1.24644-003 (2 spec.); K84-06-07. AMS 1I.25266-004 (1

spec.); K84-22-02. AMS 1.28749-004 (1 spec.); K88-17-03. AMS I.29812-001 (1 spec.); K89-15-02.
REFERENCES. — McMillan in Gomon et al. (1994); McMillan (1995).

FIGURE 114. Trachyrincus longirostris (Ginther, 1878). AMS 1.25266-004. From Kapala stn K84-22-02, off Gabo Island,
Vic., in 1052-1079 m.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 501

ACKNOWLEDGMENTS

We take this opportunity to express our deepest appreciation and thanks to the many persons and
institutions that supported in one way or another the production of this work. Tomio Iwamoto espe-
cially thanks John R. Paxton for initiating this project, and the Australian Museum for making possi-
ble a three-month’s visit to Australia through a visiting fellowship. Additional support was provided
by CSIRO Marine Research for a visit to Hobart; Martin F. Gomon (NMV) for a visit to Melbourne;
the New Zealand Foundation for Research and Technology Contract MNZ603 to Clive Roberts
(Programme Leader), Museum of New Zealand Te Papa Tongawera, for a visit to Wellington; and the
California Academy of Sciences In-House Research Fund. Yuri I. Sazonov (ZMMGU) and Yuri N.
Shcherbachev (IOAN) provided information on the type specimens of Bathygadus furvescens. Peter
McMillan (National Institute of Water and Atmosphere, Wellington, NZ) provided help and informa-
tion on a number of NSW species, especially those found also in New Zealand.

Ken Graham is appreciative of the cooperation and assistance of FRV Kapala’s captains and
crew throughout his 25 years of service on the vessel. Terry Gorman (Senior Biologist on Kapala,
1970-1988) is acknowledged for his foresight in instigating and implementing the Kapala deepwater
surveys, and is thanked for his continual encouragement.

For loans of material and general curatorial assistance, we thank Doug Hoese, John Paxton, Mark
McGrouther, Kerryn Parkinson, Sally Reader, and other AMS staff; Peter Last, David Wright,
Alastair Graham, and Gordon Yearsley (CSIRO Ian Munro Ichthyological Collection); Martin
Gomon (NMV); Clive Roberts and Andrew Stewart (NMNZ); Peter McMillan (NIWA); Jon Fong
and David Catania (CAS); Yuri I. Sazonov (ZMMGU); Yuri N. Shcherbachev (IOAN); Roland J.
McKay and Jeff Johnson (QM). Jon Fong, Mysi Hoang, and Julie Mounts (CAS) assisted with digital
images. David Pollard (NSW Fisheries) reviewed the manuscript and provided helpful suggestions
for its improvement.

502 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

APPENDIX |. Data for FRV Kapala stations cited in text. (Locations are approximate midpoints of trawls: lati-

tude °S, longitude °E; * bottom depth for midwater trawl stations.)

Station AMS Reg. No. Date Location Depth (m)
K71-05-03 15967 6-IV-71 3304 0eriN Se 52? 270-280
K71-05-04 15968 6-IV-71 33042? 4iSie50? 366-366
K71-05-06 15969 FAV 350A8. 151047: 357-366
K71-06-04 15970 15-IV-71 33-46. W552 503-503
K71-07-03 15973 DICIVETI 33°35 51 59° 375-384
K71-08-03 15974 28-IV-71 32252". 152039" 366-375
K71-08-05 15975 29-IV-71 334? AS20012 549-567
K71-09-01 15976 1-71 32°48” 152°44° 585-595
K71-11-09 15987 8-VII-71 34°59" 151-07, 366-366
Ki71-13-02 15994 30-VII-71 37°42 15001 Ss 402-408
K71-13-06 15995 2-VIMET!1 35°27’, 150°49” 549-549
K72-04-01 16565 19-IX-72 33°45’, 15 1°49" 457-457
K72-04-02 16577 19-IX-72 30380 ole 457-457
K72-05-05 16589 AEX-72 33 447. Ioie53. 549-600
K72-06-05 17859 I32X272 33°48’, 151°47’ 400-405
K72-07-01 17866 6-XE72 B85 7 SIC4 5? 729-730
K72-07-04 17867 9-X]-72 38c402 ASiles6s 729-730
K72-07-15 17316 Tox 72, 33°50. USde52° 770-780
K75-01-02 18726 2-15 33°36’, 151°59” 784-795
K75-02-08 18770 4-VI-75 B45 7 151° 10 732-805
K75-03-02 18774 1 VETS 37-42 50°13" 402-421
K75-05-02 24127 8-VIII-75 AAC iiralisd 26 402-411
K75-05-03 18838 18-VIII-75 33°04. 152033" 448-466
K75-05-04 18839 19-VIII-75 Baroy N52 05, 622-658
K75-05-05 20452 19-VIII-75 33°357 15202’ 805-841
K75-05-05 24613 19-VIII-75 33°35°. 152°02 805-841
K75-05-08 19076 21-VIII-75 SACZ0P TIS TOUS: 494-512
K75-07-03 19085 16-IX-75 3024 1520593 450-460
K76-04-03 19197 30-IV-76 33-46. 151-507 485-494
K76-05-04 19198 4-V-76 33°45" 15°51" 604-604
K76-06-03 19202 11-V-76 B32 N2 3 an52° 23" 600-604
K76-07-01 19205 26-V-76 930307 1558" 375-384
K76-23-01 19862 13-XII-76 34 7DA NSAIDS: 732-768
K76-24-03 19860 20-XII-76 33°33. 152-02: 823-823
K76-24-04 19859 21-XI-76 33°29" 1'52°06" 8232823
K77-07-10 21806 26-VI1-77 33°30’, 152°0S’ 604-604
K77-13-10 20118 23-VIII-77 29252?-.153°43" 503-512
Kiiel3-12 20301 23-VIII-77 DOES 32) 53-42: 503-503
K77-16-16 21669 QS 1NET7 33°33, 152502” 604-604
K77-18-01 20064 I6-XTT 34P 11152903" *2200-2380
K77-19-05 20071 3-XIETT 34°38". 151° 16" *2750-2930
K77-21-01 20484 DIEXIETT. 34°39 AS51020: 695-695
K77-22-03 20097 29-XI-77, 37°41", 150°18” 732-162
K77-22-06 20096 30-XI-77 37°40’, 150°20’ 823-823
K77-23-06 20485 6-XII-77 33°38. 15156: Tig=7a0
K77-23-07 20098 6-XII-77 33°32’. 152°03" 914-914
K77-23-09 21805 7 XMETT 330097 152-257 585-594
K77-23-10 20477 FeXMeTT 33°10’, 152°24’ 732-732
Kog-23ei2 20099 XI 77. 33°34, 152°01" 823-823
K77-23-13 20068 SX 7 33-267, 1527107 878-896
K77-24-03 20307 [G2 0-77, 33°21 152-267 * 1830-2750
K77-24-10 20314 14=X01-77, 337327. 1520357 *3660-3840
K77-24-11 20315 14-XII-77 34°09’, 152°01’ *2470-2560
K78-01-01 30407 22-III-78 33042 15°51" 448-457
K78-09-05 20518 2-VI-78 28°027-153-59- 549-549
K78-16-07 20435 2-VIII-78 29°47’, 153°44” 421-439

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

503

Station AMS Reg. No. Date Location Depth (m)
K78-17-07 23689 16-VIII-78 28°00’, 153°58” 411-411
K78-17-10 20459 17-VIII-78 28°01’, 154°00’ 549-549
K78-17-11 21793 17-VIII-78 98-037! 153°58° 411-411
K78-17-14 23993 17-VIII-78 DET 153053" 174-201
K78-23-08 21795 6-XI-78 28°03’, 154°04’ 732-741
K78-23-09 20651 6-XI-78 27°56’, 154°03” 549-549
K78-26-16 24037 7-XII-78 3849-15155’ 823-850
K78-27-05 23885 12-xt- 7S B4°55' 151-13" 801-827
K79-15-01 29535 2X9 33°48”, 151°49” 439-439
K79-15-03 26932 3-X-79 a8caie. 152°02” 402-402
K79-19-07 21369 28-XI-79 32°55". 153°02’ *3600-3700
K79-20-04 23710 4-XII-79 33°34’, 152°04’ P13- 7352
K79-20-06 25933 4-XII-79 33°36’, 152°06’ 914-933
K79-20-13 21722 6-XII-79 33°32’, 152°06’ 823-823
K79-20-15 21724 6-XII-79 33°37’, 152°06’ 1005-1010
K80-05-01 21725 13-V-80 33042) 151°52” 439-550
K80-20-05 40274 9-XII-80 B8u37, 152-03" 960-988
K81-17-03 24619 9-IX-81 33°46’, 151949” 439-475
K81-18-05 23862 15-IX-81 347407, 51°15" 520-530
K82-17-01 23470 122X-82 33432 151°53° 475-494
K82-24-02 23486 20-XII-82 33°47’, 151°49” 457-475
K83-06-01 24101 25-VII-83 38045 S055" 805-841
K83-06-02 24054 26-VII-83 34°36’, 151919” 869-878
K83-07-11 24100 10-VIII-83 33°04’, 152°34’ 960-997
K83-08-01 24055 18-VIII-83 33°45’, 151°59” 933-942
K83-08-02 24056 18-VIII-83 33°37’, 152°04’ 860-896
K83-09-01 24060 22-Vili-83 33047. 151°58" 942-960
K83-09-02 24059 23-Vill-83 3323075 152°10° 933-969
K83-09-04 24057 24-VIII-83 BA5532 151° 14" 951-978
K83-12-04 24157 2TAX-83 38°18’, 149948” 997-1015
K83-13-01 24150 [72X283 88,98 720152-05° 988-1015
K83-13-02 2418] 1$2X283 302577: 152°43° 960-988
K83-14-01 24172 D5K-83 35°28", 150°53” 978-1024
K83-14-02 24187 D5-X288 35°287. 150°53° 988-1024
K83-14-03 24565 D5-X183 35°28", 150°53’ 1033-1042
K83-14-05 24356 26-X-83 6 Scheu lo beat i 1042-1061
K83-14-06 24173 26-X-83 34°54’, 151914’ 1097-1116
K83-14-08 24178 DIEXeSS 38230 2152-06" 978-1006
K83-14-09 2445] JIEK=83 330321152"10’ 1042-1061
K83-15-01 24419 2x83 32°08 153°07" 910-950
K83-15-02 24462 22X1283 32°04’, 153°08’ 942-978
K83-18-01 24357 30-XI-83 94555215114" 969-1024
K83-18-02 24355 30-XI-83 FAP5520151° 16" 1105-1152
K83-19-02 24424 6-XII-83 35°29", 150°53” 1033-1070
K84-04-10 24624 11-IV-84 BATS FS 143” 1097-1134
K84-04-11 25415 11-IV-84 34°54". 151°14’ 988-1024
K84-06-03 24645 1-V-84 38 °292. °152°08” 777-823
K84-06-04 24659 1-V-84 33°29", 152°09” 914-933
K84-06-06 24625 2-V-84 335307..152°10° 1042-1106
K84-06-07 24644 2-V-84 gau0o"159°44" 1170-1198
K84-08-02 24778 IPNI=84 BAS A 151°13” 814-850
K84-08-03 24658 22-V-84 34952 151°14’ 869-924
K84-08-05 24774 23-V-84 35°38”, 150°44’ 805-850
K84-10-03 24771 17-VII-84 S252 46° 963-1039
K84-10-06 24820 18-VII-84 32°04’, 153°09” 960-969
K84-10-08 24988 19-VII-84 33r4 128152°04" 1097-1134
K84-11-07 24992 1-VIII-84 35°27 50°55" 1079-1116
K84-11-09 25273 2-VIII-84 S4P5a ISIC 1G: 1161-1207
K84-13-03 28713 24-VIII-84 36-27, 150°20° 411-457

504

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Station AMS Reg. No. Date Location Depth (m)
K84-14-01 24854 3-IX-84 FANT T DT: 439-512
K84-15-01 24850 10-IX-84 33:45 151-51 411-439
K84-15-03 24852 11-uxes4 33534 152-01" 512-530
K84-16-04 24979 25-IX-84 33°34 152-03" 722-777
K84-16-05 24991 25-IX-84 33°36, 152-03" 814-832
K84-16-13 24990 D7-1K-84 Saag Sd 58: 905-924
K84-16-14 24993 27-IX-84 33°42’, 152°04’ 1042-1070
K84-16-15 24980 DIAK-84 337432. 52°00" 960-997
K84-17-01 24989 3-X-84 84°15. 1519307 668-704
K84-17-03 24851 4-X-84 BACs Te ASIC 19: 768-786
K84-17-04 24981 4-X-84 3493672151°20° 860-878
K84-17-05 24860 4-X-84 34°48’, 151°16’ 914-969
K84-18-03 aS Oy) 10-X-84 344872151" 13° 732-750
K84-18-06 25126 11=xX-84 33°49’, 151956” 914-924
K84-19-04 25264 16-X-84 33°35’, 152°08’ 1025-1244
K84-20-03 25290 121-84 33°43’, 152°01’ 969-1006
K84-20-04 24978 -X1-84 335377 152-07" 1070-1125
K84-20-05 25095 1-X1-84 33.30% 152° 13° 1170-1207
K84-22-02 25266 PIEXI84 37°40% 150021" 1052-1079
K85-17-02 26240 14-XI-85 Chey areal oh leit 421-457
K85-17-02 26446 14-XI-85 33°36 205 1257- 421-457
K85-20-10 26221 10-XII-85 32°35". 152-49" 154-157
K85-21-04 26247 19-XII-85 33°34’, 152°09” 1024-1052
K85-21-06 25932 19-XII-85 33°43. 151°53° 439-466
K86-01-05 26394 11-11-86 33°29", 152°06’ 454-523
K86-01-06 26002 11-I]-86 83.35? 152001" 657-662
K86-01-07 26245 11-II-86 33742? 151°59° 819-889
K86-01-08 26000 12-11-86 337322, 152° 10" 951-1015
K86-01-09 26001 12-11-86 S377 8p S204. 1116-1207
K86-10-07 26453 22-IV-86 3320279152933" 439-512
K86-10-14 26756 23-IV-86 33°00’, 152°36’ 457-503
K87-02-01 26981 19-II-87 3292 153°01" 896-960
K87-14-02 26998 5-VIII-87 3471'8?= 151°30° 636-781
K87-16-02 27647 27-VIII-87 32°23’, 153°03” 878-951
K87-23-02 27609 B=xIes7 34°53. 151°C 503-658
K87-24-01 29600 7-X11-87 33352 TSS” 783-978
K87-24-02 30737 S°XTIES7 32°56’, 152°40’ 832-997
K87-24-03 28189 8 XGIES7 32°53’, 152°46’ 863-960
K87-24-04 29805 8-XII-87 32°39°* 152° 54° 887-951
K87-24-05 29738 9-XII-87 31°54. 153°12’ 479-922
K87-25-06 29741 16-XII-87 352287,.150 52. 933-960
K88-04-06 28475 23-III-88 34°55’, 151°15" 1116-1152
K88-04-08 DIG 24-III-88 33°33’, 152°08” 1024-1143
K88-04-09 28071 24-II1-88 38 9 aIS2 011° 905-969
K88-05-01 DTTNS 29-III-88 35°30’, 150°54’ 1134-1189
K88-08-04 27720 4-V-88 32°04’, 153°08” 1024-1079
K88-08-05 28372 4-V-88 32°06’, 153°09” 1070-1106
K88-08-06 28100 4-V-88 32°08’, 153°09” 1024-1079
K88-08-07 27721 5-V-88 32°56’, 152°44’ 1006-1079
K88-08-08 29601 5-V-88 354057. 152-34: 896-951
K88-08-09 TGQ? 5-V-88 33°00’, 152°38’ 704-750
K88-10-02 TNO 17-Ve88 32°50” 152°48° 1024-1061
K88-10-04 OSTIT 17-V-88 B9852". 15248" 1079-1097
K88-11-01 27638 14-VI-88 33°43". 152°07° 1024-1042
K88-12-02 27643 21-VI-88 32202 9953°10° 990-1020
K88-12-03 2959] 21-VI-88 3174679153718" 1005-1042
K88-14-04 27637 4-VIII-88 35°30’, 150°53” 988-1024
K88-16-04 29340 17-VIII-88 329567. 0152-45" 1042-1061
K88-17-03 28749 31-VIII-88 33°28", 152°14” 1143-1198

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES

Station

K88-17-06
K88-20-01
K88-20-02
K88-20-03
K88-21-02
K88-21-03
K89-03-15
K89-06-02
K89-06-04
K89-06-05
K89-07-01
K89-07-04
K89-07-05
K89-08-01
K89-08-02
K89-09-01
K89-09-03
K89-09-06
K89-09-07
K89-09-09
K89-1 1-02
K89-12-02
K89-12-03
K89-12-04
K89-12-05
K89-13-01
K89-13-02
K89-15-01
K89-15-02
K89-15-03
K89-15-04
K89-16-02
K89-17-02
K89-17-03
K89-17-04
K89-17-06
K89-17-07
K89-17-08
K89-17-09
K89-18-02
K89-18-04
K89-19-01
K89-19-02
K89-20-01
K97-01-21
K97-01-22
K97-02-01

AMS Reg. No. Date Location Depth (m)
28900 1-IX-88 5329 SIS” 1033-1070
29809 8-XI-88 B2°547.15 2°47" 1070-1097
29297 8-XI-88 32°50 152°48" 1024-1097
29298 8-X1-88 327508152748" 988-1033
29747 3=XdIE88 32°48"..1/52°477 841-933
29746 3-XII-88 32°40’, 152°50’ 713-750
29385 27-11-89 38°06’, 149°42’ 152-159
29799 11-IV-89 2756). 152 47: 1042-1061
29808 11-IV-89 32°51, 152-48" 1090-1134
29813 12-IV-89 che ose pa 722-768
29803 18-IV-89 35°05 151-07" 695-768
32431 19-IV-89 35°42?" 150°43" 887-960
29807 19-IV-89 35°41", 150°43’ 1015-1042
29811 9-V-89 33°43’, 151959” 805-869
29801 10-V-89 330282 S2716 1134-1189
29804 16-V-89 33704’..152°36' 896-960
28988 16-V-89 32051°s 152,49" 1024-1061
29806 17-V-89 33°307..152-07° 732-796
29605 18-V-89 33°39’, 152°05’ 1024-1088
29825 18-V-89 33°27". 152710" 741-768
28070 8-VI-89 32°517) 152°492 1006-1052
30738 14-VI-89 32°51, 152°48’ 1079-1143
29762 14-VI-89 32°43’, 152°49” 713-796
29798 15-VI-89 32°06’, 153°10’ 1033-1079
29797 15-VI-89 32°03’, 153°09” 914-997
29823 29-VI-89 32 °3 38152. 59° 896-969
28477 30-VI-89 3344” 152203” 1116-1170
29827 3-VIII-89 32 567 152-45" 1024-1061
29812 3-VIII-89 32-51 152 49° 1061-1097
29757 3-VIII-89 32°51? 15248" 933-988
29756 4-VIII-89 3393972 151959" 677-750
30394 10-VIII-89 BAP ATE, 1S 1218? 1134-1225
29749 15-VIII-89 32°30 S27 52? 814-850
29754 15-VIII-89 32°30’, 153°00’ 1006-1052
29750 16-VIII-89 B1P4T7.. WS 3°48? 1024-1052
29753 16-VIII-89 3527, 153° 16° 878-933
29742 17-VIII-89 3°07. 153.09" 1079-1143
29752 17-VIII-89 32°06’, 153°09” 1024-1061
30304 17-VIII-89 B21? 15306: 823-860
29745 22-VIII-89 94°56 15 1°15’ 1090-1143
39052 22-VIII-89 34°45”, 151°16’ 950-990
29737 31-VIII-89 35°29", 150°55’ 1116-1134
29761 31-VIII-89 35°29’, 150°53” 1024-1061
29743 7-IX-89 3304 al 52-007 805-869
39957 29-IV-97 3743 al S000" 219-227
39958 30-IV-97 37239 4 150-172 543-567
38576 27-V-97 35°33", 150°46’ 505-549

506 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

LITERATURE CITED

ALcock, A. 1894. Natural history notes from H. M. Indian Marine Survey Steamer ‘Investigator,’ Commander
C. E. Oldham. R. N., commanding. Series 2, No. 11. An account of a recent collection of bathybial fishes
from the Bay of Bengal and from the Laccadive Sea. J. Asiat. Soc. Bengal 63 (pt. 2)(2):115—137, pls. 6-7.

ANDREW, N. L., K. J. GRAHAM, K. E. HODGSON, AND G. N. G. GORDON. 1997. Changes after twenty years in rel-
ative abundance and size composition of commercial fishes caught during fishery independent surveys on
SEF trawl grounds. NSW Fisheries Final Report Series No. 1. 210 pp.

ARAI, T. 1979. Additional information on a rare macrourid fish, Mesobius antipodum, from New Zealand. Japan.
J. Ichthyol. 25(4):286—289, figs. 1-3, table 1.

ARAI, T. AND T. IWAMOTO. 1979. A new species of the macrourid fish genus Coelorinchus from off Tasmania,
New Zealand, and the Falkland Islands. Japan. J. Ichthyol. 26(3):238—246.

ARAI, T. AND P. J. MCMILLAN. 1982. A new macrourid fish, Coelorinchus biclinozonalis from New Zealand, and
redescription of C. australis from Australia. Japan. J. Ichthyol. 29(2):115—126.

BARNARD, K. H. 1925. Descriptions of new species of marine fishes from S. Africa. Ann. Mag. Nat. Hist., ser. 9,
15(87):498-504.

DOLLo, L. 1909. Cynomacrurus Piriei, Poisson abyssal nouveau recueilli par |’Expédition Antarctique
Nationale Ecossaise. Note préliminaire. Proc. Roy. Soc. Edinburgh 29(4):3 16-326.

FILHOL, H. 1884. Explorations sous-marines—Voyage du “Talisman.” Nature, Paris (558):182—186.

GARMAN, S. 1899. Report on an exploration off the west coasts of Mexico, Central and South America, and off
the Galapagos Islands, in charge of Alexander Agassiz, by the US Fish Commission steamer “Albatross”
during 1891, XXVI (The Fishes). Mem. Mus. Comp. Zool. Harvard Coll. 24:1-431.

GILBERT, C. H. 1905. The deep-sea fishes of the Hawaiian Islands. Pp. 575-713, figs. 230-276, pls. 66-101 in
The aquatic resources of the Hawaiian Islands, D. S. Jordan, and B. W. Evermann, eds., Bull. U. S. Fish
Comm. 1903, 22(pt. 2, sect. 2).

GILBERT, C. H. AND F. CRAMER. 1897. Report on the fishes dredged in deep water near the Hawaiian Islands, with
descriptions and figures of twenty-three new species. Proc. U. S. Natl. Mus. 19:403-435.

GILBERT, C. H. AND C. L. HuBBS. 1920. The macrourid fishes of the Philippine Islands and the East Indies. U. S.
Natl. Mus. Bull. 100, 1 (pt. 7):369-588, figs. 1-40.

GOMON, M. F., J. C. M. GLOVER, AND R. H. KuiTER, eds. 1994. The fishes of Australia’s south coast. State Print,
Adelaide. 992 pp.

GON, O. AND P. C. HEEMSTRA, eds. 1990. Fishes of the Southern Ocean. J. L. B. Smith Institute of Ichthyology,
Grahamstown. 462 pp., 12 pl.

GoopE, G. B. AND T. H. BEAN. 1883. Reports on the results of dredging under the supervision of Alexander
Agassiz, on the east coast of the U.S. XIX. Report on the fishes. Bull. Mus. Comp. Zool. Harvard
10(5):183-226.

GORMAN, T. B. AND K. J. GRAHAM. 1975. Deepwater prawn survey off New South Wales. Pp. 162—172 in Pro-
ceedings of First Aust. Nat. Prawn Sem., P. C. Young, ed. Australian Government Publishing Service, Can-
berra.

GRAHAM, K. J. 1990. Report for Cruises 89-06 to 89-20 on the NSW mid-slope between Crowdy Head and
Batemans Bay during April-September, 1989. Kapala Cruise Report 107. NSW Fisheries, Cronulla, Austra-
lia. 22 pp.

GRAHAM, K. J., N. L. ANDREW, AND K. E. HODGSON. 2001. Changes in relative abundance of sharks and rays on
Australian South East Fishery trawl grounds after twenty years of fishing. New Zealand J. Mar. Freshwater
Res. 52:549-561.

GRAHAM, K. J. AND T. B. GORMAN. 1985. New South Wales deepwater prawn fishery research and development.
Pp. 231-243 in Second Aust. Nat. Prawn Sem., P. C. Rothlisberg, B. J. Hill and D. J. Staples, eds. NPS2,
Cleveland , Australia.

GRAHAM, K. J.,G. W. LIGGINS, AND J. WILDFORSTER. 1996. NSW continental shelf trawl survey results for Year
2: 1994. Kapala Cruise Report No. 115. NSW Fisheries, Cronulla, Australia. 63 pp.

GRAHAM, K. J.,G. W. LIGGINS, J. WILDFORSTER, AND B. Woop. 1995. NSW continental shelf trawl-fish sur-
vey results for Year 1: 1993. Kapala Cruise Report No. 114. NSW Fisheries, Cronulla, Australia. 52 pp.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 507

GRAHAM, K. J., B. R WOOD, AND N. L. ANDREW. 1997. The 1996-97 survey of New South Wales upper slope
trawling grounds between Sydney and Gabo island. Kapala Cruise Report No. 117. NSW Fisheries, Cro-
nulla, Australia. 96 pp.

GUNTHER, A. 1877. Preliminary notes on new fishes collected in Japan during the expedition of H. M. S. “Chal-
lenger.” Ann. Mag. Nat. Hist., ser. 4, 20:433-447.

. 1878. Preliminary notices of deep-sea fishes collected during the voyage of H. M. S. “Challenger.” Ann.

Mag. Nat. Hist., ser 5, 2:17—28.

. 1887. Report on the deep-sea fishes collected by H. M. S. Challenger during the years 1873-76. Rep.
Sci. Res. Challenger 22(pt. 57):1-Ixv + 1-268, pls. 1-73.

HAEDRICH, R.L. AND P.T. POLLONI. 1976. A contribution to the life history of a small rattail fish,
Coryphaenoides carapinus. Bull. So. Calif. Acad. Sci. 75(2):203-211.

Howes, G. J. 1989. Phylogenetic relationships of macrouroid and gadoid fishes based on cranial myology and
arthrology. Pp. 113—128 in Papers on the systematics of gadiform fishes, D. M. Cohen, ed. Nat. Hist. Mus.
Los Angeles County, Sci. Ser. 32. 262 pp.

Howes, G. J. AND O. A. CRIMMEN. 1990. A review of the Bathygadidae (Teleostei: Gadiformes). Bull. Br. Mus.
Nat. Hist. (Zool.) 56(2): 155-203.

Huss, C. L. AND T. IWAMOTO. 1977. A new genus (Mesobius), and three new bathypelagic species of
Macrouridae (Pisces, Gadiformes) from the Pacific Ocean. Proc. Calif. Acad. Sci. ser. 4, 41(7):233-251.

IwAMOTO, T. 1970. The R/V Pillsbury Deep-Sea Biological Expedition to the Gulf of Guinea, 1964-65. 19.
Macrourid fishes of the Gulf of Guinea. Stud. Trop. Oceanogr. (4)(pt.2):3 16-431.

. 1974. Nezumia (Kuronezumia) bubonis, anew subgenus and species of grenadier (Macrouridae: Pisces)

from Hawaii and the western North Atlantic. Proc. Calif. Acad. Sci. ser. 4, 39(22):507—516.

. 1982. Ventrifossa johnboborum, anew grenadier from the western Pacific (Macrouridae: Pisces). Austr.

Zool. 21(pt.1):55-61.

. T. 1986. Family No. 93: Macrouridae. Pp. 330-341 in Smiths’ sea fishes, M. M Smith. and P. C.

Heemstra, eds. Macmillan South Africa, Johannesburg.

. 1990. Macrouridae. Pp. 90-317 in FAO Species Catalogue, vol. 10. Gadiform fishes of the world. An
annotated and illustrated catalogue of cods, hakes, grenadiers and other gadiform fishes known to date,
D. M. Cohen, T. Inada, T. Iwamoto, and N. Scialabba. FAO, Rome.

IWAMOTO, T. AND M. E. ANDERSON. 1994. Review of the grenadiers (Teleostei: Gadiformes) of southern Africa,
with descriptions of four new species. Ichthyol. Bull. J. L. B. Smith Inst. Ichthyol. (61):1—28.

IWAMOTO, T. AND P. MCMILLAN. 1997. A new grenadier, genus 7rachonurus, from New Zealand and Australia
(Macrouridae, Gadiformes, Pisces). Mem. Mus. Victoria 56(pt. 1):255—259.

IWAMOTO, T., P. MCMILLAN, AND Y. N. SHCHERBACHEV. 1999. A new grenadier, genus Caelorinchus, from
Australia and New Zealand (Pisces, Gadiformes, Macrouridae). New Zealand J. Mar. Freshwater Res.
33(1):49-S4.

IWAMOTO, T. AND N. R. MERRETT. 1997. Pisces Gadiformes: Taxonomy of grenadiers of the New Caledonian re-
gion, southwest Pacific. Pp. 473-570 in Résultats des Campagnes MUSORSTOM, vol. 18, A. Crosnier, ed.
Mem. Mus. Natn. Hist. Nat. 176.

IWAMOTO, T. AND Y. I. SAZONOV. 1988. A review of the southeastern Pacific Coryphaenoides (sensu lato) (Pis-
ces, Gadiformes, Macrouridae). Proc. Calif. Acad. Sci. 45(3):35—82, figs. 1-9.

IWAMOTO, T. AND Y.N. SHCHERBACHEV. 1991. Macrourid fishes of the subgenus Chalinura, genus
Coryphaenoides, from the Indian Ocean. Proc. Calif. Acad. Sci. 47(7):207-233, figs. 1-17, tables 1—7.

IWAMOTO, T. AND A. WILLIAMS. 1999. Grenadiers (Pisces, Gadiformes) from the continental slope of western
and northwestern Australia. Proc. Calif. Acad. Sci. 51(3):105—243.

JORDAN, D. S. AND C. H. GILBERT. 1904. Macrouridae. Pp. 602-621 in List of fishes dredged by the steamer A/-
batross off the coast of Japan in the summer of 1900, with descriptions of new species and a review of the
Japanese Macrouridae, D. S. Jordan and E. C. Starks. Bull. U. S. Fish Comm. 22(1902):577—630, pls. 1-8.

Last, P. R., E. O. G. SCOTT, AND F. H. TALBOT. 1983. Fishes of Tasmania. Tasmanian Fishery Development
Authority, Hobart. 563 pp.

LAST, P. R. AND J. D. STEVENS. 1994. Sharks and rays of Australia. CSIRO Australia. 513 pp., 84 pls.

LEVITON, A. E. AND R. H. Gibss, JR. 1988. Standards in herpetology and ichthyology. Standard symbolic codes
for institution resource collections in herpetology and ichthyology. Supplement No. |: additions and correc-
tions. Copeia 1988(1):282.

508 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 21

Leviton, A. E.,R. H. GipBs, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology:
Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.
Copeia 1985(3):802-832.

LowE, R. T. 1843. Notices of fishes newly observed or discovered in Madeira during the years 1840, 1841 and
1842. Proc. Zool. Soc. Lond. 11-91.

MAKUSHOK, M. 1976. The new rattail Coryphaenoides subserrulatus sp. n. (Macrouridae, Osteichthys) from the
area south of New Zealand. Pp. 144—55 in Biology and distribution of tropical deep-sea fishes, N. V Parin,,.
ed. Trudy Inst. Okeanol. Akad. Nauk SSSR, Moscow, v. 104:1—226. [In Russian. ]

MARSHALL, N. B. 1964. Bathypelagic macrourid fishes. Copeia 1964(1):86—93.

. 1973. Family Macrouridae. Pp. 496-665 in Fishes of the western North Atlantic, D. M. Cohen, ed.
Mem. Sears Found. Mar. Res. (1)(pt.6).

MARSHALL, N. B. AND T. IWAMOTO. 1973. Genus Coryphaenoides. Pp. 565—600 in Fishes of the western North
Atlantic, D. M. Cohen, ed. Mem. Sears Found. Mar. Res. (1)(pt.6).

MCCANN, C. AND D. G. MCKNIGHT. 1980. The marine fauna of New Zealand: Macrourid fishes (Pisces: Gadida).
New Zealand Oceanogr. Inst. Mem. 61:1—91.

MCCULLOCH, A. R. 1907. The results of deep sea investigations in the Tasman Sea. II. The expedition of the Woy
Woy. Fishes and crustaceans from eight hundred fathoms. Rec. Aust. Mus. 6:345-355, 6 pls.

MCCULLOCH, A. R. 1926. Report on some fishes obtained by the F.I.S. “Endeavour” on the coasts of
Queensland, New South Wales, Victoria, Tasmania, South and South-western Australia. Part V. Biological
results of the fishing experiments carried on by the F. I. S. “Endeavour” 1909-1914, 4(pt. 5):157—216, pls.
43-56.

MCMILLAN, P. J. 1995. Review of trachyrincine grenadier fishes (Pisces: Macrouridae) from New Zealand, with
a description of a new species of Trachyrincus. New Zealand J. Mar. Freshw. Res. 29:83-91.

. 1999. New grenadier fishes of the genus Coryphaenoides (Pisces: Macrouridae); one from off New Zea-
land and one widespread in the southern Indo-West Pacific and Atlantic Ocean. New Zealand J. Mar.
Freshw. Res. 33:481—-489.

MCMILLAN, P. J. ANDC. D. PAULIN. 1993. Descriptions of nine new species of rattails of the genus Caelorinchus
(Pisces, Macrouridae) from New Zealand. Copeia 1993(3):8 19-840.

MERRETT, N. R. AND T. IWAMOTO. 2000. Macrourid fishes of the New Caledonia region, Southwest Pacific
Ocean; taxonomy and distribution, with ecological notes. Résultats des Campagnes MUSORSTOM, vol.
21. Mém. Mus. Natn. Hist. Nat. 184:723-781.

Munro, I. S. R. 1957. Handbook of Australian Fishes. No. 14. (Aust.) Fish. Newsletter 16 (8).

NoRMAN, J. R. 1939. Fishes. Scientific Report of the John Murray Expedition. British Museum (N.H.), London,
7(1): 1-116.

OKAMURA, O. 1970. Fauna Japonica. Macrourina (Pisces). Academic Press, Tokyo. 216 pp., 64 pls.

. 1982. [Macrouridae] /n Fishes of the Kyushu-Palau Ridge and Tosa Bay, O. Okamura, K. Amaoka, and

F. Mitani, eds. Japan Fish Resource Conserv. Assoc., Tokyo. 435 pp.

. 1984. [Macrouridae] /n Fishes of the Okinawa Trough and the adjacent waters. I. The intensive research

of unexploited fishery resources on continental slopes, O. Okamura and T. Kitajima, eds. Japan Fish. Re-

source Conserv. Assoc., Tokyo. 414 pp.

. 1989. Relationships of the suborder Macrouroidei and related groups, with comments on Merlucciidae
and Steindachneria. Pp. 129-142 in Papers on the systematics of gadiform fishes, D. M. Cohen, ed. Nat.
Hist. Mus. Los Angeles County, Sci. Ser. 32. 262 pp.

PAXTON, J. R., D. F. HOESE, G. R. ALLEN, AND J. E. HANLEY. 1989. Zoological catalogue of Australia. Vol. 7,
Pisces, Petromyzontidae to Carangidae. Austr. Govt. Publ. Serv., Canberra. 664 pp.

RADCLIFFE, L. 1912. Descriptions of a new family, two new genera, and twenty-nine new species of anacanthine
fishes from the Philippine Islands and contiguous waters. Proc. U. S. Natl. Mus. 43:105—140, pls. 22-31.

RICHARDSON, J. 1839. [. . . account of an interesting collection of fish formed at Port Arthur in Van Diemen’s
Land .. .]. Proc. Zool. Soc. London. 7:95—100.

. 1846. Ichthyology of the voyage of H. M. S. Erebus and Terror under the command of Captain Sir James
Clark Ross, R.N., F.R.S., during the years 1839-1843. E. W. Janson, London. 139 pp, 60 pls.

SAZONOV, Y. I. 1981. /diolophorhynchus andriashevi gen. et sp. n. (Osteichthyes, Macrouridae) from the Aus-
tralia-New Zealand region. Zoologicheskiy Zhurnal 60, vol. 9:1357—1363.

SAZONOV, Y. I. AND T. IWAMOTO. 1992. Grenadiers (Pisces, Gadiformes) of the Nazca and Sala y Gomez ridges,
southeastern Pacific. Proc. Calif. Acad. Sci. 48(2):27—95, 37 figs., 7 tables.

IWAMOTO AND GRAHAM: GRENADIERS OF NEW SOUTH WALES 509

SAZONOV, Y. 1. AND Y. N. SHCHERBACHEV. 1982a. On the taxonomic position and distribution of Coelorinchus
matamua (McCann and McKnight)(Gadiformes, Macrouridae). Pp. 42-47 in Insufficiently studied fishes
of the open ocean, N. V. Parin, ed. Inst. Okeanol. Akad. Nauk. SSSR, Moscow. 140 pp. [In Russian. ]

. 1982b. A preliminary review of grenadiers related to the genus Cetonurus Giinther (Gadiformes,

Macrouridae). Descriptions of new taxa related to the genera Cetonurus Giinther and Kumba Marshall. [In

Russian, with English summary.] Vopr. Ikhtiol. 22 (5):707—721, figs. 1-4. [also Engl. transl., J. Ichthyol.

22(5):1—15]

. 1985. Preliminary review of grenadiers of the Cetonurus group (Gadiformes, Macrouridae). II. The ge-
nus Cetonurus Ginther: taxonomic characters of the group. J. Ichthyol. 25(3):12—26, figs. 1-2, tables 1-2.

SHCHERBACHEV, Y. N. 1987. Preliminary list of thalassobathyal fishes of the tropical and subtropical waters of
the Indian Ocean. Vopr. Ikhtiol. (1):3-11. [In Russian.] [English version in J. Ichthyol. 27 (2):3746.]

SHCHERBACHEV, Y. N. AND T. IWAMOTO. 1995. Indian Ocean grenadiers of the subgenus Coryphaenoides, genus
Coryphaenoides (Macrouridae, Gadiformes, Pisces). Proc. Calif. Acad. Sci. 48(14):285—3 14, figs. 1-8, ta-
bles 1-3.

SHCHERBACHEV, Y. N., Y. I. SAZONOV, AND T. IWAMOTO. 1992. Synopsis of the grenadier genus Kuronezumia
(Pisces: Gadiformes: Macrouridae), with description of a new species. Proc. Calif. Acad. Sci. 48(3):97—108,
figs. 1-9, table 1.

SHCHERBACHEV, Y. N., Y. I. SAZONOV, AND A. S. PIOTROVSKIY. 1979. On the discovery of Trachonurus villosus
and species of the genus Mesobius (Macrouridae, Osteichthyes) in the Indian Ocean. J. Ichthyol.
19(1):16-23, figs. 1-2, tables 1-2.

SMITH, H. M., AND L. RADCLIFFE. 1912. [See RADCLIFFE, 1912]

TiLzey, R. D. J. 1994. Introduction. Pp 15—40 in The South East Fishery, R. D. J. Tilzey, ed. Bureau of Resource
Sciences, Canberra, Australia.

TRUNOV, I. A. 1981 (1980). [A new genus and species of grenadier, Haplomacrourus nudirostris (Osteichthyes,
Macrouridae)]. Vopr. Ikhtiol. (1980). t. 20,. 1(120):3-11. [Also published in J. Ichthyol. tome 20,
1(120):3-11.]

TRUNOV, I. A. 1983. On the characteristics of some species of the genus Coelorinchus (Macrouridae) from the
southeastern Atlantic Ocean. Report 1. Vopr. Ikhtiol. 23(6):894—904. [In Russian. ]

WEBER, M. 1913. Die Fische der SIBOGA-Expedition. Siboga Exped. 57:1—719, pls. 1-12.

WEBER, M. AND L. F. DE BEAUFORT. 1929. The fishes of the Indo-Australian Archipelago, vol. 5. E. J. Brill,
Leiden. 458 pp.

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Volume 52, No. 22, pp. 511-548, 5 figs. |

NOV 2 6 2001

| November 13, 2001
'

The Octandrous and Dodecandrous Species of Topobea
(Melastomataceae) in Mexico and Central America

by

Frank Almeda
Department of Botany, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

A regional revision of the octandrous and dodecandrous species of Topobea is presented
that recognizes 24 species in Mexico and Central America. This summary includes a key,
descriptions, distributional and phenological information, discussions of useful taxonomic
characters, and a brief review of what is known about the reproductive biology of Topobea.
Rationale is given for placing nine species in synonymy. Illustrations are provided for four
new species (7. amplifolia, T. dimorphophylla, and T. gerardoana from Costa Rica; T.
tetramera from Panama), and for 7. multiflora, a species erroneously reported for Costa
Rica as T. calycularis. Representative specimens are cited to document the geographic dis-
tribution and range of variability for each species.

RESUMEN

Se presenta una revisién de las 24 especies de Topobea en México y America Central que
tienen ocho y doce estambres. Este resumen incluye una clave, descripciones, informacion
sobre distribucién y fenologia, y discusiones de caracteres taxonomicos muy utiles en
Topobea. También se presentan ilustraciones para cuatro especies nuevas (7. amplifolia, T.
dimorphophylla, y T. gerardoana de Costa Rica; T. tetramera de Panama), y para T.
multiflora, una especie erréneamente reportada para Costa Rica como 7. calycularis.
Especimenes representativos son citados para documentar la distribucion geografica y el
rango de variabilidad de cada especie.

Topobea, with approximately 70 species, is one of two genera comprising the neotropical tribe
Blakeeae. This tribe is readily recognized by its prevailingly 6-merous axillary flowers that are indi-
vidually subtended by two pairs of decussate bracts, baccate fruits, and ovoid to pyramidal seeds with
a smooth testa (Almeda 1990).

Most species of Topobea are shrubby epiphytes of wet forest habitats; some are terrestrial shrubs,
and a few attain arborescent dimensions. More than 75% of the species occur in a narrow equatorial
band centered in Colombia, ranging south to Ecuador and north to Costa Rica. Another dozen or so
species extend north to southern Mexico and south to Bolivia and Brazil. Because of their showy flow-
ers, the terrestrial species, some of which are locally common, have been collected with increasing
frequency in recent decades. Because so many species of Topobea are obligate epiphytes with very lo-
cal distributions, newly discovered species have come to light with increasing frequency as the forest
canopies of tropical America have received increased exploratory attention in the last two decades.

Topobea was last treated in its entirety by Cogniaux (1891) who recognized 24 species, five of
which were reported for the Mesoamerican region. Subsequent regional floras (Gleason 1958; Stand-
ley 1924, 1938; Standley and Williams 1963; Winkler 1965) collectively attributed 17 species of

51]

512 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

Topobea to Mexico and Central America, only six of which are recognized here as distinct taxa. This
study of the octandrous and dodecandrous species of Mexico and Central America and my recent revi-
sion of the Central American hexandrous clade (Almeda 2000a) recognize a total of 29 species of
Topobea for the region. Including the four new species proposed here, over half of these have been de-
scribed or transferred to Topobea in the last 17 years.

TAXONOMIC TREATMENT
Topobea Aubl., Hist. Pl. Guiane Fr. 1:476. 1775.
TYPE: Topobea parasitica Aubl.

Trees, shrubs, or woody vines, often epiphytic with glabrous or variously pubescent terete to
quadrate branchlets. Leaves coriaceous or chartaceous, 3—7-nerved or plinerved, often with the trans-
verse secondary veins closely spaced, straight and parallel. Flowers 6-merous (rarely 4-merous), dip-
lostemonous (haplostemonous in five Mesoamerican species), axillary, solitary or fascicled, typically
pedunculate in the upper leaf axils and subtended by two pairs of decussate, free or partially fused,
coriaceous or foliaceous bracts inserted at the base of the hypanthium. Hypanthium campanulate or
suburceolate; calyx persistent, truncate or 6-lobate (4-lobate in one species). Petals white or pink, ma-
genta or some combination of these colors, glabrous to sparsely pubescent abaxially, sometimes
ciliolate at the margins. Stamens 12 or 6 (8 in one species), isomorphic and glabrous; anthers lin-
ear-oblong or subulate, uniporose or biporose with 2 dorsally-inclined apical pores that are approxi-
mate, cleft and divergent, or often confluent at anthesis; connective simple and unappendaged or
thickened and modified basally at or near the filament insertion into a spur or caudiform appendage.
Stigma punctiform to capitate. Ovary completely or partly inferior but varying to superior in a few
species, usually 6-locular (consistently 2- or 4-locular in some species). Fruit baccate; seeds clavate to
cuneate or narrowly pyriform.

Until recently, the tribe Blakeeae was thought to be constant with respect to ovary position and
number of perianth parts and ovary locules (Almeda 1990, 2000a). Descriptive literature on the tribe
describes it as having 6-merous flowers with 12 stamens (dodecandrous) and a 6-locular inferior
ovary (Almeda 1990). This characterization still holds true for Blakea, which is distinguished from
Topobea by its laterally compressed anthers that are biporose, oval, oblong, or elliptic and obtuse to
rounded apically with 2 well-separated (and typically minute) apical pores (Almeda 2000b).
Topobea, on the other hand has uniporose or biporose anthers that are linear-oblong to ob-
long-subulate (usually not compressed laterally) with dorsally-inclined apical pores that are com-
monly confluent or cleft and divergent. Most species of Topobea are similar to Blakea in having
6-merous flowers that are dodecandrous and a 6-locular ovary. This appears to be the plesiomorphic
condition in the tribe. Critical study of the Mesoamerican species of Topobea reveals that only five
species have completely inferior ovaries and a surprising number exhibit an evolutionary reduction
series involving stamen and ovary locule number. The species of Topobea treated here fall into three
groups based on stamen number. The most extraordinary one in this regard is T. fetramera. As its spe-
cific epithet implies it has 4-merous flowers that are octandrous (with eight stamens) and its ovary 1s
inferior and 4-locular. The second, a hightly derived group of five species, is characterized by
6-merous flowers that are hexandrous (with 6 stamens) and an inferior ovary that is either 2- or
4-locular (Almeda 2000a). The third group consists of the remaining 23 dodecandrous species. Sev-
enteen of these have a 6-locular ovary, five are consistently 4-locular, and one, 7. albertieae, is
prevailingly 4-locular with the occasional 3- or 5-locular ovary. Reduction in stamen number is con-
sistently correlated with an inferior ovary position in the hexandrous and octandrous species. This
kind of character correlation, however, breaks down among the dodecandrous species. The following

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA

PLATE |. Topobea fragrantissima Almeda. Original gouache and acrylic on hotpress illustration board by Meg Stalcup in
July, 2001.

514 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

examples serve to illustrate this point: 7. aeruginosa has 6:12:6 (petals: stamens: locules) with a com-
pletely inferior ovary; T. brenesii also has 6:12:6 but its ovary is wholly superior; 7. watsonii with
6:12:6 has an ovary that is 1/3-inferior; 7. dodsonorum has 6:12:4 and a 1/2-inferior ovary; T. pittieri
has 6:12:4 and a 3/4-inferior. Among the dodecandrous species, the lack of correlation between ovary
position and locule number provides information for identifying phenetic gaps and useful character
combinations for species delimitation. It provides a challenge, however, when attempting to deter-
mine derived character states for phylogenetic reconstruction. The general assumption among stu-
dents of the Melastomataceae has been that a superior ovary is plesiomorphic and an inferior ovary is
apomorphic. The diversity in ovary position among the species of Topobea considered here is sugges-
tive of homoplasy and leads to some intriguing questions about the forces driving this kind of charac-
ter evolution.

Another character of taxonomic importance in Topobea is the presence of foliar domatia. Nine of
the twenty-nine Mesoamerican species of Topobea produce specialized pit, pocket, or hair tuft cham-
bers (acarodomatia) in the vein axils of some or all abaxial leaf surfaces. The mite-leaf domatium as-
sociation is generally a mutually beneficial relationship (Walter and Proctor 1999). The domatia
provide shelter and protection for eggs and moulting mites. The mites evidently benefit the plants by
feeding on fungal spores and the eggs of predaceous insects.

In addition to ovary position and meristic differences in petal, stamen, and ovary locules, a num-
ber of other characters are important for the delimitation and identification of Topobea species. These
include size, shape, and degree of fusion of floral bracts, presence or absence of lateral anther sac fu-
sion, modifications of the anther connective into appendages, and details of the anther pores. Staminal
material is essential for generic placement and for definitive identification of many species. It has
been necessary to use these characters in the key that follows despite the fact that all of them are not al-
ways available on every specimen. Because of this I have also tried, where possible, to include vegeta-
tive characters which are more readily accessible.

Information on the reproductive biology of Topobea is available for only two species, 7. brenesii
and 7. maurofernandeziana (including T. durandiana as reported in Lumer 2000). Both of these spe-
cies are endemic to Mexico and/or Central America. According to Lumer (2000), these two species
are self-compatible and capable of producing viable seeds without outcrossing. Lumer found that ex-
perimentally selfed flowers of 7. brenesii produced significantly more seeds than outcrossed flowers.
The showy flowers of both species offer pollen as the primary reward which attracts several species of
bees despite interspecific flower differences in size and texture. The bee pollinators of Topobea use
the same buzzing method of pollen collecting that is widespread among the melastomes and other
flowering plants with poricidal anthers. Bees alight on the flowers, bend their bodies over the anther
cluster, and vibrate their indirect flight muscles in a way that results in rapid evacuation of pollen from
the terminal anther pores of a flower. The ejected pollen is deposited on the bee’s ventral side and
readily transferred to the stigma of the next flower visited (Lumer 2000). Because both species of
Topobea studied are self-compatible, insect-mediated selfing may constitute a significant factor in ef-
fecting optimal pollination and seed set.

KEY TO THE OCTANDROUS AND DODECANDROUS SPECIES OF TOPOBEA

1. Flowers 4-merous and diplostemonous (with eight stamens per flower); ovary 4-locular.......... T. tetramera
1’. Flowers 6-merous and diplostemonous (with twelve stamens per flower); ovary 4-locular or 6-locular.
2. Outer floral bracts conspicuously decurrent on and imparting a winged aspect to the floral peduncle T. mcphersonii
2'. Outer floral bracts not conspicuously decurrent on the floral peduncle.
3. Both surfaces of mature leaf blades moderately to copiously covered with smooth ferrugineous hairs
0.5—3(—9) mm long.
4. Mature leaves of a pair markedly unequal in size with the larger blade commonly six to twelve times the
size of the smaller blade; floral peduncle 0.8—1.4 cm long; inner floral bracts fused basally for 3.5—5 mm
to form a bowl-like collar; petals 0.5—0.6 0.5 cm; anthers 1.5-3 mmlong ...... T. dimorphophylla

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA StS

4'. Mature leaves of a pair somewhat unequal in size with the larger blade typically not more than two or three times
the size of the smaller blade; floral peduncle 3.5—5.2 cm long; inner floral bracts free to the base; petals 1.6—-2.3
4S e7icm:tanthers—7- Sin ON eee nat es ee eee, eS ete ee ee ee) Seas T. intricata

3’. Both surfaces of mature leaf blades not covered with ferrugineous smooth hairs.

5. Some or all mature leaf blades typically bearing domatia (pit, pocket, or hair tuft domatia) on the abaxial surface
in the basal angles between the median vein and each of the two proximal lateral veins.
6. Anther sacs laterally connate for half or more of their length.

7. Mature leaf blades bearing hair tuft domatia on the abaxial surface in the angles between the median vein
and each of the two proximal veins; each anther with two confluent apical pores; connective
dorso-basally apendaged; ovary 6-locular.

8. Uppermost cauline nodes covered with caudate-acuminate stipuliform flaps ca. 3 mm long that
envelop caducous tufts of hairs, the distal portions of each flap + caducous with age and leaving a
prominent interpetiolar ridge or corky line; outer floral bracts fused at the base for 4-6 mm; anthers

G=Simmillong Rage Sen Cees Sees She oR ae eek BNR. T. multiflora
8’. Uppermost cauline nodes not as above; outer floral bracts free from one another; anthers 4-5 mm long
GET REPE VERE 244 ey ot 2k ee ale RAY ost sige er eearey Sine e! Beeb ho T. pluvialis

7'. Mature leaf blades bearing perforated pit domatia on the abaxial surface in the angles between the median
vein and each of the two proximal lateral veins; each anther with two separate apical pores; anther
connective simple and unappendaged; ovary 4-locular .................... T. parvifolia

6’. Anther sacs completely free from one another.

9. Some or all abaxial foliar surfaces bearing hair tuft domatia that are sparsely to moderately covered with
barbellate hairs in the angles formed between the median vein and the proximal pair of lateral veins.

10. Anthers 2.5 mm long, the surface granulose along the lower ventral half of the thecae; each anther
with a solitary apical pore; connective prolonged dorso-basally into a deflexed caudiform append-
APE TOVALVAOIOCUIARANS Sere Meme #6 tend ceetre Teies Een e Deay. ware cere Pe T. lentii

10’. Anthers 5—7 mm long, the surface smooth throughout the length of the thecae; each anther with two
confluent apical pores; connective simple and unappendaged; ovary 4-locular. ... . T. calycularis

9'. Some or all abaxial foliar surfaces bearing pit or pocket domatia that lack hairs of any kind.

11. Calyx consisting of a truncate flange or broadly flattened into low undulations; each anther
LeLMminAalin on twOICONLUCHt PORES) mews ee Meteo wa net aera Me owes cole T. laevigata

11’. Calyx consisting of well-defined lobes; each anther terminating in a solitary pore. Principal leaves
markedly dimorphic in size at each node, the larger blades broadly rounded and + subpeltate at the
WETS esau A a 0° 35 Ge cS? bho A eae en Ba es eee eee a ok le eter T. dodsonorum
12’. Principal leaves not markedly dimorphic in size at each node, all mature blades acute to obtuse

at the base.
13. Outer floral bracts 4.5—7.5 x 3-4 mm; petals 12-15 mm long; anther connective
unappendapedzovary O-loculate. tue ees cans cee a) cet cae ee T. suaveolens
13’. Outer floral bracts 1.2-2.5 x 1.5—2.5 mm; petals 7-8.5 mm long; anther connective
prolonged dorso-basally into a + horizontal toothlike appendage 0.5 mm long; ovary
@-loculaniwytys 28 crt eu en ee ity Seater tele de rehtale ea ra Shay: T. pittieri
5'. Mature leaf blades lacking domatia on the abaxial surface in the basal angles between the median vein and each of
the two proximal lateral veins.
14. Calyx lobes 13-17 mm long, covered with a dense indument of spreading barbellate or plumose hairs.

15. Mature leaf blades dentate at least distally; anthers laterally connate for much of their length, each anther

sac with 2 confluent, dorsally-inclined pores; ovary apex modified into a glandular-puberulent fluted

cConesandicollar7—9immyl one etree aaa seek “anes oath face cee apcuces op oeetis T. calophylla
15’. Mature leaf blades entire; anthers free from one another, each anther with 2 divergent, dorsally-inclined
apical pores; ovary apex glabrous and not elevated into acone and stylarcollar ...... T. aeruginosa

14’. Calyx truncate and flangelike or with tnangular or depressed-triangular lobes 1-5 mm long, glabrous or if
pubescent then the indument not as above.
16. Floral bracts and calyx lobes strongly undulate-recurved apically; inner floral bracts fused basally for
7-8 mm; each anther with 2 divergent, dorsally-inclined apical pores ............. T. brenesii
16’. Floral bracts and calyx lobes not strongly undulate-recurved apically; inner floral bracts free; each anther
sac with 2 confluent, dorsally-inclined apical pores (if anther pores are divergent then the anther
connectives are unappendaged).
17. Blades of the principal leaves broadly rounded to rounded-emarginate at the apex; abaxial foliar
surfaces consistently but inconspicuously glandular-punctate .............. T. albertieae
17'. Blades of the principal leaves typically acuminate but sometimes varying to acute or caudate at the
apex; abaxial foliar surfaces not glandular-punctate.

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Volume 52, No. 22

18. Anthers of a flower all free from one another.
19. Anthers releasing pollen by a solitary apical pore.

20. All mature leaf blades glabrous throughout, the transverse secondary veins on the abaxial surface spaced
mostly 0.25 mm apart at the widest portion of the blade; ovary 4-locular........... T. fragrantissima

20’. Some mature leaf blades sparingly covered with a caducous indument of rusty brown subulate and conic hairs
on the abaxial surface, the transverse secondary veins on the abaxial surface spaced 1-3 mm apart at the
widest portion of the blade; ovary 6-locular ... ~~. --- 1+ see eee eee ee eee T. gerardoana

19’. Each anther releasing pollen by two confluent apical pores.
21. Calyx consisting of well-defined lobes.
22. Mature leaf blades 21-25.3 x 11-14 cm; floral bracts glabrous, anther connective prolonged dorso-basally
into a deflexed appendage; ovary apex elaborated into a sleeve-like distally lobulate collar 2-3 mm long
ne are ere ee or ee eer eer odes B bl oresoed-oro Grin Op a og oo c T. amplifolia
22'. Mature leaf blades 9.5—15 x 5-10 cm; floral bracts moderately to sparsely covered with an indument of
caducous matted hairs; anther connective unappendaged; ovary apex lacking acollar.... . T. standleyi
21’. Calyx consisting of a truncate flange or the lobes broadly flattened into low undulations .. . . . T. laevigata
18’. Anthers of a flower laterally connate for half or more of their length.
23. Anthers conspicuously granulose along at least the lower ventral half of the thecae; transverse secondary veins on
the abaxial surface of mature leaf blades spaced 0.25—0.5 mm apart at the widest portion of the blade . 7. watsonii
23’. Anthers smooth throughout; transverse secondary veins on the abaxial surface of mature leaf blades spaced

1-4 mm apart at the widest portion of the blade.

24. Calyx tube (fused portion of calyx below the free lobes) 6-8 mm long; filaments 7-9 mm long; anther thecae
linear-oblong; ovary apex covered with a caducous ring of minute glandular hairs surrounding the stylar scar
cold, Cease dee Dhesel folese tier okeitoerioal oe. GTi eu Als ed eee, Ot eee, T. maurofernandeziana

24'. Calyx tube 2-4 mm long; filaments 9-12 mm long; anther thecae subulate; ovary apex elevated into a cone
about 2 mm long and a shallow collar about 0.5—0.75 mm long that may disappear as the ovary enlarges on
maturedruits., s 2tidy steed. c cee Qacteem ae ators i orlh GIGeeeS bo SPRR none Reems T. parasitica

1. Topobea aeruginosa (Standl.) L. O. Williams, Fieldiana Bot. 31:35. 1965.

Blakea aeruginosa Standl., Field Mus. Nat. Hist., Bot. Ser. 17:381. 1938.

TyPE. — HONDURAS. Comayagua: near the summit of the ridge above El Achote, 1850 m, |
Aug. 1936, Yuncker et al. 6267 (holotype: F!; isotypes: K!, NY!, US!).

Tree (or epiphytic shrub, fide Yuncker) 3—10 m tall. Upper cauline internodes bluntly quadrate.
Uppermost internodes, vegetative buds, petioles, peduncles, and floral bracts densely covered with a
ferrugineous indument of barbed or plumose hairs. Leaves coriaceous when dry, 10—23.5 x
4.5—17 cm, elliptic-oblong to elliptic-ovate, the apex short-acuminate, the base obtuse to rounded, the
margin entire, S—7-nerved, the outermost pair of primaries often concealed by revolute foliar margins
when dry, the blade adaxially glabrous and glossy, abaxially covered with barbed or plumose hairs.
Flowers solitary or paired in axils of upper branchlets; peduncles 1.7—3 cm long. Outer floral bracts
1.5-2.6 x 1-1.5 cm, free or fused basally for 3-5 mm, lanceolate, the apex long-attenuate; inner
bracts 1.5—1.8 x 1 cm, free, ovate, the apex long-acuminate. Calyx lobes linear-oblong, |.3—1.5 cm
long. Petals 6, glabrous but caducously ciliate, |.5—1.7 x 0.5—0.8 cm, pink, obovate, the apex rounded.
Stamens 12; filaments 0.7—0.8 cm long; anthers free, 0.9-1.1 cm long, | mm wide, yellow, lin-
ear-oblong, each with 2 divergent, dorsally-inclined apical pores; connective elevated dorso-basally
into a blunt oblong appendage 0.5—1 mm long. Ovary completely inferior, 6-locular, glabrous and not
elevated into a cone or stylar collar. Style glabrous, 1.5—1.8 cm long; stigma punctiform. Mature berry
1.3—1.5 x 1-1.3 cm. Seeds | mm long, brown, cuneate to narrowly pyriform.

DISTRIBUTION AND PHENOLOGY. — Local in cloud forests of Honduras and Nicaragua at
1200-1500 m. Flowering and young fruiting specimens have been collected from July through De-
cember.

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA ey

REPRESENTATIVE SPECIMENS EXAMINED. — NICARAGUA. Matagalpa: camino a Aranjuez a
menos de | km de carretera Matagalpa-Jinotega, 13°02'N, 85°55’W, 2 Jul. 1980, Moreno 1028
(CAS); along road to La Fundadora, Cordillera Central, 22 Feb. 1963, Williams et al. 24886 (F).

DISCUSSION. — This species is apparently known only from the type in Honduras. It is readily
recognized by its rusty brown indument of plumose hairs on vegetative buds, abaxial leaf surfaces,
floral peduncles, floral bracts and calyx lobes. Its free anthers are also distinctive in having two diver-
gent apical pores.

2. Topobea albertiae Wurdack, Phytologia 55:146. 1984.

TYPE. — COLOMBIA. Antioquia: Fincas Montepinar and Las Palmas, Vereda Quebrada Larga,
municipio Guatapé at the line with municipio San Rafael, elev. 1800 m, 4 Sep. 1982, Albert de
Escobar et al. 2278 (holotype: HUA; isotype US!)

Epiphytic shrub 2.5—3 m tall or reportedly a tree 5—12 m tall. Uppermost branchlets quadrate be-
coming rounded with age, the young vegetative buds and uppermost nodes covered with caducous
simple hairs. Nodes on older branches becoming notably thickened with elevated interpetiolar lines or
ridges. Mature leaves of a pair essentially equal or only somewhat unequal in size, glabrous adaxially
but inconspicuously glandular-punctate abaxially; petioles 1-3.8cm long; blades coriaceous,
5.7-16(-19) x 3.7—9.2(-12) cm, obovate to elliptic-obovate to elliptic, the apex rounded to
rounded-emarginate, the base broadly acute to obtuse, the margin entire and sometimes revolute when
dry, 5-nerved, the transverse secondary veins spaced 1—2.5 mm apart at the widest portion of the
blade. Flowers erect, borne in clusters of 24 in each leaf axil of uppermost branches; peduncles
1.1-1.5 cm long, commonly lenticellate distally. Floral bracts erect, concave and closely enveloping
the hypanthium, essentially glabrous or sparsely and caducously covered with a mixture of appressed
hairs 0.5—1 mm long and amorphous branlike hairs, the margin often fimbriate-ciliolate; outer bracts
8—10 x 10 mm, fused basally for 3-5 mm, + oblong to oblate, with a mostly rounded-truncate apex
that sometimes varies to bluntly acute; inner bracts 7-10 10 mm, free but imbricate, ovate to oblate,
the apex + truncate to broadly rounded. Hypanthium (at anthesis) cupulate, 4-5 x 5 mm. Calyx tube
5—6 mm long, erect and + cylindric; calyx truncate and without evident lobes. Petals 6, glabrous and
reflexed, 1.8-2.8 x 1.4-1.9cm, pale pink or white flushed with pink, the claw typically white,
obovate, the apex + truncate-rounded, the margin caducously ciliolate. Stamens 12, the filaments
1.4-1.7 cm long, declinate, complanate, glabrous; anthers laterally connate for about half their length
and forming a subparallel horizontal platform, 10—13 = 0.75—1 mm, yellow, oblong-subulate, each
with 2 confluent, dorsally-inclined apical pores; connective thickened dorsally and prolonged
dorso-basally into an acute appendage 2—2.5 mm long. Ovary superior or 1/2- to 1/3-inferior,
(3—)4(-5)-locular, elevated into a cone 4-5 mm high but lacking a collar. Style glabrous 1.7—2 cm
long, stigma punctiform. Mature berries and seeds not seen.

DISTRIBUTION AND PHENOLOGY. — Local in cloud forests of central Panama to Colombia at
100-2300 m. Collected in flower in January, February, July, and from October through December, in
fruit from January through April and October through November.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Cocle: area of El Valle, 2 km E of La
Mesa, N slope of Cerro Gaital, 8°38’N, 80°7’W, 16 Nov. 1983, Churchill 3870 (CAS). Darién:
Serrania del Darién, just below Cerro Mali, Gentry et al. 16841 (DUKE). Panama: near Cerro Jefe
about 0.5—2 miles beyond road junction on the dirt road to Alto Pacora, 9°15'N, 79°30’W, 11 Jan.
1989, Almeda et al. 6153 (CAS, PMA).

DISCUSSION. — This species is distinctive in having glandular-punctate abaxial leaf surfaces, a
cylindric calyx tube that lacks well-defined calyx lobes, and anther thecae that are laterally connate for
a portion of their length. The flowers are also extraordinary in producing a clove-like fragrance remi-

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niscent of that encountered in some neotropical orchids that attract euglossine bees. Topobea
albertiae is unusually variable in characters which are typically constant and diagnostic among its
congeners. For example, the number of ovary locules in most specimens examined is four but some
flowers have only three locules and others have as many as five. The degree of ovary fusion to the
hypanthial wall is equally puzzling because the ovary is completely superior in some flowers exam-
ined and 1/2 to 1/3 inferior in other flowers.

3. Topobea amplifolia Almeda, sp. nov. (Fig. 1)

TYPE. — COSTA RICA. Limon: Canton de Talamanca. Bratsi, Amubri, Alto Lari, Kivut.
Afluente innominado del Rio Lari, margen izquierda, 9°23'25”N, 83°04'25”W, 1200 m, 21 Mar.
1992, Herrera 5407 (holotype, CAS!; isotypes, CR, INB!, MO!).

Arbor 6 m alta. Petioli 3—5 cm longi, lamina 21—25.3 x 11—14 cm elliptica vel elliptico-ovata
apice acuminata vel acuta basi acuta vel obtusa, 5—7-nervata ad maturitatem chartacea et glabra,
nervis secundariis 0.25 mm inter se distantibus. Flores 6-meri in quoque nodo superiore 4—S,
pedunculis | .3—2 cm longis, bracteae exteriores 0.8—1.2 x 0.7—0.9 cm ovatae vel elliptico-lanceolatae
ca. 3 mm coalitae apice truncato-rotundata plerumque acuto vel cuspidato; bracteae interiores 0.9 x
1 cm omnino liberae. Hypanthium (ad torum) 0.8—0.9 longum; calycis tubus 2— 3 mm longum, lobis
3-4 mm longis. Petala alba, 1.9 x 1.3 cm oblongo-ovata vel obovata. Filamenta 6 mm longa; antherae
6 x 2 mm inter se non cohaerentes, dorsaliter biporosae; connectivum ad basim dorsaliter | mm
descendenti armatum. Stylus 1.5 cm; ovarium 6-loculare et 2/3 inferum, collo 2-3 mm alto glabro.

Reportedly a tree 6 m tall. Uppermost branchlets with thickened interpetiolar ridges at each node,
the internodes rounded-quadrate becoming rounded with age. Vegetative and very young floral buds
copiously but caducously covered with an amorphous rusty brown scurfy indument. Mature leaves of
a pair essentially equal or only slightly unequal in size, glabrous throughout; petioles 3—S cm long;
blades chartaceous, 21—25.3 x 11—14 cm, elliptic-ovate to elliptic, the apex acuminate to abruptly
acute, the base acute to obtuse, the margin inconspicuously crenulate, 5—7-nerved, the outermost
intramarginal pair commonly depressed and inconspicuous; the transverse secondary veins spaced
0.25 mm apart at the widest portion of the blade. Flowers erect to spreading, borne in clusters of four
or five in each leaf axil of distal branches; peduncles |.3—2 cm long. Floral bracts glabrous, closely en-
veloping the hypanthium; outer bracts 0.8—1.2 x 0.7—0.9 cm, fused basally for 3 mm, ovate varying to
elliptic-lanceolate, the apex abruptly acute to cuspidate; inner bracts 0.9 x | cm, free, depressed-ovate
to suborbicular, the apex commonly retuse and often mucronate. Hypanthium (at anthesis)
campanulate, 0.8—0.9 x 0.8—-0.9 cm. Calyx tube 2—3 mm long; calyx lobes (fruiting hypanthium),
erect, 3-4 mm long and 3-4 mm wide basally, oblong-ovate, rounded apically and covered with
callose-thickened teeth at the abaxial apex. Petals 6, glabrous, 1.9 x 1.3 cm, white, obovate, the apex
rounded, entire. Stamens | 2; filaments complanate and glabrous, 6 mm long; anthers free, 6 x 2 mm,
reportedly yellow-brown, subulate, each with 2 confluent, dorsally-inclined apical pores; connective
thickened dorsally and prolonged dorso-basally into a deflexed tooth-like appendage | x 0.25 mm.
Ovary 2/3 inferior, 6-locular, elaborated apically into a glabrous, sleeve-like distally lobulate collar
2—3 mm long. Style glabrous, 1.5 cm long; stigma + clavate and obliquely flattened when dry. Imma-
ture berry 0.8 x 1 cm. Seeds cuneate to narrowly obovoid, | mm long, brownish with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Known only from low-elevation rainforest in southeastern
Costa Rica near the Panamanian border at 1200 m. The type and only known collection, which was
made in March, is in flower and young fruit.

DISCUSSION. — The distinctive features of 7. amplifolia include the closely spaced (0.25 mm)
transverse secondary veins on abaxial foliar surfaces (Fig. 1B), callose-thickened teeth at the abaxial

ALMEDA: 7OPOBEA IN MEXICO AND CENTRAL AMERICA S19

FicurE 1. Topobea amplifolia Almeda. A. habit, ca. 1/5; B. representative leaf (abaxial surface, ca. 1/3; C. young fruiting
hypanthium with decussate floral bracts, ca. 3; D. petals (adaxial surface), 2; E. stamens, dorsal view (left) and profile view
(right), 5; F. stylar collar, style, and stigma, 3;G. seeds, 15. (A-G from Herrera 5407.)

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apex of each calyx lobe (Fig. 1 C), dorso-basally appendiculate anther connectives (Fig. 1 E), and dis-
tally lobulate collar that envelops the style base (Fig. 1F). Among Mesoamerican species, it most re-
sembles 7. multiflora that differs in having hair tuft domatia on abaxial foliar surfaces, stipuliform
nodal flaps, pink petals, and anther thecae that are laterally connate for at least half of their length.
Topobea amplifolia is also superficially similar to 7. superba Naudin of Colombia and T.
subscaberula Triana of Colombia and Ecuador. The former differs by its nodal stipuliform flaps
(1-2 mm long), hair tuft acarodomatia with roughened hairs 1—-2.5 mm long where primary leaf veins
diverge from one another abaxially, secondary transverse veins spaced |.5—4 mm apart, and anther
thecae that are laterally coherent for about half of their length. Topobea subscaberula differs from T.
amplifolia in having young leaves and pedicels that are moderately but caducously puberulous with
pinoid hairs 0.1—0.2 mm long, primary leaf vein divergence (abaxial) covered with roughened hairs
about 1 mm long, secondary transverse veins spaced 2—2.5 mm apart, apically obtuse outer floral
bracts, and an ovary apex that is elevated into a broad crateriform dome but not elaborated into a
lobulate collar.

ETYMOLOGY. — The epithet amplifolia is derived from the Latin words amplus, ample or large,
and folius, -leaved, in reference to the large leaves of this species.

4. Topobea brenesii Standl., Field Mus. Nat. Hist., Bot. Ser. 18:842. 1938.

TYPE. — COSTA RICA. Alajuela: La Palma de San Ramon, 1250 m, 13 Mar. 1929, Brenes 6732
(holotype: F!; isotypes: CR!, NY!).

Epiphytic shrub 2-4 m tall. Uppermost branchlets rounded-quadrate becoming rounded with
age, the uppermost internodes, young vegetative buds, pedicels, floral peduncles, floral bracts (espe-
cially abaxial surfaces), hypanthia, and calyx lobes copiously covered with a caducous mixture of
elongate roughened hairs and less conspicuous stellulate or branlike hairs, the upper cauline nodes co-
piously setose with mostly smooth or sparingly roughened hairs. Mature leaves of a pair equal to
somewhat unequal in size; petioles 0.2—0.7 cm long; blades coriaceous, 4.5—14 x 2.1—6.7 cm, obovate
to elliptic, the apex rounded to obtuse or acute, the base broadly rounded to truncate, margin entire,
blade 3-nerved or if 3-plinerved then the inner pair of primary veins diverging from the median vein
0.6—1 cm above the blade base (on abaxial surface), the transverse secondary veins spaced |.5—3 mm
apart at the widest portion of the blade, the blade adaxially covered with a sparse cover of elongate
roughened and branlike hairs but commonly glabrous at maturity, abaxially sparsely covered with
elongate roughened and branlike hairs especially on the elevated primary veins. Flowers erect, | to 3
in each leaf axil of distal branches; peduncles 0.2—1.3 cm long. Floral bracts closely enveloping
hypanthium and strongly undulate-recurved apically; outer bracts 8-9 x 6.5—7 mm, free or fused ba-
sally for 0.5—1 mm, broadly oblong; inner bracts 9-10 x 13-14 mm, fused basally for 7-8 mm, semi-
circular. Hypanthium (at anthesis) campanulate, 7-8 x 7-8 mm. Calyx tube 3-4 mm long, erect and
cupulate to campanulate; calyx lobes 4 x 4-6 mm, oblong and rounded at the recurved apex. Petals 6,
glabrous, I|—1.5 x 0.5—1.4 cm, pale pink, obovate. Stamens 12; filaments 4-6 mm long, declinate and
glabrous; anthers free, 4.5—5.5 x 1 mm, yellow, each opening by 2 divergent, dorsally-inclined pores;
connective thickened dorsally and prolonged dorso-basally into a horizontally divergent or recurved
knobby appendage 0.5 mm long. Ovary wholly superior, 6-locular, elevated at the glabrous apex into
a lobulate rimlike collar that forms a wide circle around the stylar scar. Style glabrous, 1.2—1.4 cm
long; stigma punctiform. Berry 1—1.2 x 1—1.2 cm. Seeds narrowly ovoid to cuneate, 1—1.25 mm long,
beige with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Endemic to Costa Rica where it is local and uncommon in
cloud forests of the Cordillera de Tilaran and adjacent slopes southeast to the San Ramon region and
east to the western slopes of Volcan Viejo in the Cordillera Central at 950-1560 m. Collected in

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 52]

flower from January through April, in fruit from January through April, July, August and probably in-
tervening months.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela: Canton de San Ramon.
Los Angeles, Colonia Palmarena. Cuenca media de Rio San Lorenzo, camino a la mina de yesa,
10°12’50”N, 84°35'15”W, 20 Feb. 1991, Herrera et al. 4918 (CAS, CR, INB, MO): Canton de San
Carlos, Zapote, 31 Oct. 1938, A. Smith 1306 (CAS, NY). Puntarenas: Cordillera de Tilaran. Selec-
tively logged pasture known locally as the Bull Pen about 0.5—1 km downslope from Monteverde
Cloud Forest Reserve Station, 29 Feb. 1992, Almeda & Daniel 7185 (CAS, CR).

DISCUSSION. — This species is known only from the Cordillera de Tilaran south and east to the
San Ramon and Volcan Viejo regions of Costa Rica. It is easily recognized by its completely superior
ovary, leaf blades that are rounded to truncate basally, conspicuously appendiculate anther connec-
tives, and undulate-recurved floral bracts and calyx lobes.

5. Topobea calophylla Almeda, Proc. Calif. Acad. Sci. 43:281. 1984.

TYPE. — PANAMA. Veraguas: 5 mi. W of Santa Fé on road past Escuela Agricola Alto Piedra
on Pacific side of divide, elev. 800-1200 m, 18 Mar. 1973, Croat 23000 (holotype: CAS!; isotype:
MEXU!, MO!, US!).

Coarse epiphytic shrub. Upper branches rounded to subquadrate. Distal internodes, vegetative
buds, peduncles, and floral bracts covered with a hirsute indument of ferrugineous barbellate hairs
mostly 3—9 mm long. Mature leaves of a pair somewhat unequal in size; blades firmly chartaceous to
coriaceous, 14.5—37.5 x 8.6—17.8 cm, elliptic-ovate, 5—7-nerved, the elevated transverse secondary
veins spaced 4—7 mm apart at the widest portion of the blade, adaxially glabrous, abaxially moder-
ately hirsute with barbellate hairs mostly 1—3 mm long, the apex abruptly caudate-acuminate, the base
rounded to subcordate, the margin inconspicuously dentate. Flowers erect to widely spreading, paired
or borne in clusters of three to four in leafy axils of distal branches; peduncles 2.8—4 cm long. Floral
bracts foliaceous, entire, 2—5-nerved, free, each pair closely subtending one another or separated on
the peduncle by a distance of 34 mm. Outer bracts 1.7—2.3 « 1.5—1.7 cm, elliptic-ovate, the apex
acuminate; inner bracts 1.6—1.9 x 1.3—1.7 cm, elliptic-ovate, the apex acute to acuminate. Calyx lobes
lance-triangular, 14—17 x 5-6 mm. Petals 6, glabrous, entire but sparingly glandular-ciliate, 2 x 1 cm,
reportedly pink, + spatulate, the apex acute to obtuse. Stamens 12, filaments 5 x 1.5 mm, strongly
declinate; anthers laterally connate for a good portion of their length, 8 x 1.5 mm, yellow (?), lin-
ear-subulate, each with 2 confluent, dorsally-inclined apical pores; connective thickened dorsally
near the filament insertion into a blunt callosity. Ovary completely inferior, 6-locular, apex prolonged
into a glandular-puberulent fluted cone and stylar collar mostly 7—9 mm long. Style glabrous, | 1—14 x
1 mm; stigma capitellate to truncate, the actual surface appearing somewhat crateriform. Mature
berry not seen.

DISTRIBUTION AND PHENOLOGY. — Locally common in rainforests and cloud forests from the
vicinity of Fortuna Dam to the Santa Fé region of Veraguas disjunct to Comarca de San Blas in
north-central Panama at 10—1200 m. Collected in flower from February through August, in fruit dur-
ing March and July.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Bocas del Toro: vicinity of Fortuna
Dam, below pass on Chiriqui Grande road, 8°45'N, 82°15'W, 27 June 1986, McPherson 9714 (CAS,
MO, PMA). Comarca de San Blas: Cerro Brewster, 9°18’N, 79°16'W, 21 Apr. 1985, de Nevers et al.
5384 (CAS, MO): Rio Taindi (Taimdi of maps), 6 km above confluence with Rio Mandinga, 9°25'N,
79° 11'W,5 Apr. 1986, de Nevers & Herrera 7638 (CAS, MO, PMA). Veraguas: Distrito de Santa Fe,
alrededores del Rio Primer brazo de Ulaba, 8°33’'N, 81°07'W, 6 Jul. 1996, Galdames et al. 3145

522 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

(CAS, SCZ): Boca de Concepcion, in Golfo de los Mosquitos, forest near river, 8°50’'N, 81°00'W, 6
Aug. 1987, McPherson 11394 (CAS, MO, PMA).

DISCUSSION. — This large-leaved Panamanian endemic has an unusual indument of long,
spreading, barbellate hairs on distal internodes, young buds, peduncles, and floral bracts. This to-
gether with the laterally connate anther thecae, and glandular-puberulent fluted ovary cone and stylar
collar at the ovary apex make it a standout among Mesoamerican species. For an illustration of this
species see Almeda (1984:280).

6. Topobea calycularis Naudin, Amn. Sci. Nat. Bot. 3, 18:149. 1852.

Type. — MEXICO. Chiapas: Zuluzuchiapas, Linden 650 (holotype: P!, fragment at BR!;
isotype: K!).

Epiphytic shrub or tree 2-13 m tall. Uppermost branchlets quadrate and carinate on the angles be-
coming rounded with age, essentially glabrous throughout, the young floral bract margins, floral
buds, and calyx rim sometimes covered with a caducous furfuraceous indument of stellulate or matted
fimbriate branlike hairs. Mature leaves at a node somewhat unequal in size (the larger sometimes
twice the size of the opposing one); petioles 1.3—4 cm long; blades coriaceous, 4.5—17.2 x 2.1-10 cm,
elliptic to elliptic-obovate, the apex abruptly acuminate, the base acute, the margin entire,
3—5-plinerved with the innermost primaries diverging from the median vein 0.4—1.5 cm above the
blade base (on abaxial surface) with hair tuft acarodomatia that are sparsely to moderately covered
with robust shaggy hairs (0.25—0.5 mm long) in the angles formed with the median vein, the trans-
verse secondary veins spaced 0.5—1 mm apart at the widest portion of the blade. Flowers erect, soli-
tary or in clusters of 2 to 4 in each leaf axil of distal branches; peduncles 4-9 mm long. Floral bracts +
concave and closely enveloping the hypanthium; outer bracts 3-5 x 4-5 mm, fused basally for
1.5—2 mm, rounded-triangular to ovate; inner bracts 3.54 x 4—5 mm, free but partly imbricate, semi-
circular. Hypanthium (at anthesis) campanulate, 4.5 x 4.5 mm. Calyx tube 2-3 mm long, erect and
cupulate, calyx broadly flattened into low apiculate undulations 0.5 x 2 mm. Petals 6, glabrous, 1—1.3
0.30.6 cm, white, narrowly obovate, the margin fimbriate. Stamens 12; filaments 6-7 mm long,
declinate and glabrous; anthers free, 5—7 x 0.75 mm, yellow, each with 2 confluent, dorsally-inclined
pores at the apex; connective thickened dorsally, unappendaged or with a minute dorso-basal callos-
ity. Ovary 1/3 inferior, 4-locular, apex glabrous, stylar scar evident but not elevated into a prominent
cone or stylar collar. Style glabrous, |—1.5 cm long; stigma punctiform. Berry 0.9-1.1 = 0.8—-1 cm.
Seeds narrowly pyriform to narrowly ovoid, | mm long, beige with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Often common in lowland and montane rainforests from
Chiapas, Mexico east to Guatemala (Izabal) at 320-1625 m. Collected in flower from January
through May and in November and December, in fruit from February through August, December and
probably most intervening months.

REPRESENTATIVE SPECIMENS EXAMINED. — GUATEMALA. Alta Verapaz: Chapultepec
Farm, 62 km beyond Coban on Sebol road, 20 May 1964, Contreras 4730 (CAS, LL); Pansamala,
Feb. 1887, von Tuerckheim 1135 (DS, NY, US). Izabal: Municipio El Estor La Cumbre, al NE del
Estor, 17 Jul. 1988, Tenorio et. al. 14516 (CAS, MEXU). MEXICO. Chiapas: Municipio La
Trinitaria, Lagos de Montebello National Park, 19 Nov. 1980, Breedlove & Almeda 47572 (CAS,
MEXU); Municipio of Ocosingo 70 km SW of Palenque on road to Ocosingo along the Jol Uk’ um,
Breedlove & Almeda 48335 (CAS, MEXU); Municipio of Peltalcingo, slope of Ahk’ ulbal Nab above
Peltalcingo, Breedlove 50462 (CAS, MEXU): Municipio of Independencia, 45-50 km E of Lagos de
Montebello National Park on road to Ixcan from Santa Elena, Breedlove & Almeda 57731 (CAS,
MEXU); Municipio de Tila, pie del Cerro Acavaina, Ton 7347 (CAS, MEXU); Municipio
Margaritas, Col. Maravilla, Tenejapa, Ton 9084 (CAS, MEXU).

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 523

DISCUSSION. — Topobea calycularis is distinguished by its hair tuft foliar domatia, calyx lobes
that consist of low apiculate undulations, essentially unappendaged anther connectives, and its
4-locular ovary. Only one collection examined, Tenorio et al. 14516, CAS, lacks good development
of hair tuft domatia. Topobea laevigata is the only other species with which T. calycularis might be
confused. In 7: /aevigata, the domatia, when produced, are slitlike pit domatia at the abaxial margin of
the leaf blade near the petiole-laminar junction, and the ovary is consistently 6-locular.

For Costa Rican specimens erroneously identified as 7. calycularis, see the discussion under T.
multiflora.

7. Topobea dimorphophylla Almeda, sp. nov. (Fig. 2)

TYPE. — COSTA RICA. Heredia: along Rio Peje about 0.5 km SW of back end of Vargas prop-
erty; approximately in the area where an imaginary line drawn between Magsasay (colonia penal) and
Puerto Viejo de Sarapiqui would cross the Rio Peje, 20 Feb. 1982, Hammel 11217 (holotype: CAS!;
isotypes: CR!, DUKE!, INB!, MO!, US!).

Frutex hemiepiphyticus. Ramuli sicut pedunculi folia inflorescentia hypanthiaque _pilis
1.5—3(—9) mm longis induti. Folia in quoque pari dimorpha papyracea distanterque denticulata
3—5-plinervata. Folia maiora: lamina (5.5—)9-15.5 x 2—7.5 cm elliptica vel elliptico-ovata apice
caudato-acuminata basi rotundata. Folia minora: lamina 0.9—1.7 x 0.6—1 cm ovata vel subcordata
apice caudoto-acuminata basi cordata. Flores 6-meri in quoque nodo superiori singuli, pedunculis
0.8—0.9(—1.4) cm longis; bracteae exteriores omnino liberae 0.5—0.7 x 0.3—-0.5 cm elliptico-ovata
apice acuto; bracteae interiores 0.5—0.6 x 0.4-0.5 cm ovatae apice acuta ca. 3.5—5 mm coalitae.
Calycis tubus 0.5—0.75 mm longus, lobis 0.7—0.9 x 0.2—0.3 cm. Petala 5-6 x 5 mm obovata vel
subrotundata. Antherarum thecae 1.5—3 x 0.50.75 mm inter se non cohaerentes, dorsaliter
biporosae; connectivum dorsaliter supra thecarum basim tuberculatum. Ovarium 6-loculare et
onmino inferum apice glabro (cono et collo non evoluto).

Secondary hemiepiphytic shrub with main stems growing vinelike up trunks of host trees and
secondary branches either drooping or horizontally spreading to 1-3 m long. Uppermost branchlets
mostly terete, the older branches covered with numerous short root-like protuberances (adventitious
roots?). Cauline internodes, leaf blades (both surfaces), peduncles, floral bracts, hypanthia, and calyx
lobes copiously hirsute with rusty brown hairs mostly 1.5—-3(—9) mm long. Mature leaves of a pair
markedly unequal in size; blades coarsely papery when dry, the larger one at each node (5.5—)9—15.5 x
2-7.5 cm, elliptic to elliptic-ovate, the apex caudate-acuminate, the base broadly rounded, the margin
denticulate (sometimes remotely so), 3-plinerved with an additional ill-defined intramarginal pair, the
innermost pair of primary veins diverging from the median vein 2-3 mm above the blade base, the +
transverse secondary veins spaced 2—5 mm apart at the widest portion of the blade on the abaxial sur-
face; petiole 5-17 mm long; the smaller blade 0.9-1.7 x 0.6—1 cm, ovate to subcordate, apex short
caudate-acuminate, base cordate, margin entire, 3-nerved, the transverse secondary veins not evident
on the abaxial surface; petiole barely prolonged or up to 2 mm long. Flowers erect, solitary in each axil
of uppermost leaves; peduncles 0.8—0.9(—1.4) cm long. Floral bracts green and entire; outer bracts
0.5—0.7 x 0.30.5 cm, free, elliptic-ovate, the apex acute; inner bracts fused basally for 3.5—-5 mm to
form a bowl-like collar, the free lobes broadly ovate to deltoid, 0.5—0.6 x 0.4-0.5 cm. Hypanthium at
anthesis 4—5 mm long to the torus and 4—5 mm in diameter. Calyx tube 0.5—0.75 mm long, + erect at
anthesis. Calyx lobes (on young fruit) deltoid at base but abruptly tapered to narrow linear upright seg-
ments 0.7—0.9 cm long and 0.2—0.3 cm wide at the base between sinuses. Petals 6, glabrous, 5—6
5 mm, translucent white, thin and translucent when dry, broadly obovate to subrotund, the apex +
rounded, the base shortly clawed, entire. Stamens 12, free and isomorphic; filaments |.5—-3 mm long,

524 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

{VE
SEES |
Ue aa * BESS
JE
Wes phe
ye aS
(Mlle CU

WATS:

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Wa
/ \
/
1
se si <
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NS
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FIGURE 2. Topobea dimorphophylla Almeda. A. habit, 2/5; B. foliar dimorphism ata node, 2/3; C. outer floral bract, 5; D.
inner floral bracts, 4; E. hypanthium (floral bracts removed). 3; F. petal (adaxial surface), ca. 5; G. stamens, profile view (left)
and dorsal view (right) 12;H.séeds, 15.(A from Gomez etal. 21127; B from Grayum et al. 7932; C-G from Hammel 11217;H
from Hammel & Trainer 13241.)

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 525

glabrous; anthers 1.5—3 = 0.5—0.75 mm, yellow, oblong, each with 2 confluent, dorsally-inclined
pores at the apex; connective thickened dorsally and barely elevated into a blunt callose knob
dorso-basally. Ovary completely inferior, 6-locular, glabrous at the apex which is barely elevated at
the stylar scar. Style glabrous, declinate, 9-10 mm long; stigma punctiform. Berry red at maturity,
9-10 x 9-10 mm. Seeds | mm long, brown, cuneate to narrowly pyriform, testa smooth.

DISTRIBUTION AND PHENOLOGY. — Local and uncommon in the Caribbean slopes and lowlands
of Costa Rica from the Puerto Viejo region south to the Cordillera de Talamanca (Limon) at
100—1000 m. Collected in flower and fruit from October through March and in June and July.

PARATYPES. — COSTA RICA. Alajuela: Reserva Bioldgica Monteverde Rio Pefas Blancas,
Laguna y Quebrada Celeste, 10°20'N, 84°41'W, 850 m, 4 Nov. 1989, Bello 1459 (CAS); Reserva
Forestal de Arenal, Quebrada San Gerardo, Rio Cano Negro, 10°23'N, 84°48'W, 800 m, 18 Feb.
1990, Bello 1909 (MO). Cartago: Valle Escondido, 720 m, 31 Mar. 1966, Schnell 641 (CR, US).
Heredia: Parque Nacional Braulio Carrillo, La Virgen, Sarapiqui, Sendero Transecto, 10°16’N,
84°05'W, 700 m, 11 Dec. 1988, Ballestero 53 (CAS, CR, INB). Parque Nacional Braulio Carrillo,
Fila Carrillo — Sendero La Botella, 400-800 m, 15 Feb. 1984, Gomez et al. 21127 (CAS, CR); Parque
Nacional Braulio Carrillo, forest between Rio Peje and Rio Sardinalito on Atlantic slope of Volcan
Barva, 10°17.5’N, 84°05’W, 700-800 m, 14 Nov. 1986, Grayum & Herrera 7895 (CAS, CR); Finca
La Selva, OTS Field Station on the Rio Puerto Viejo just E of its junction with the Rio Sarapiqui
100 m, 18 Jul. 1982, Hammel & Trainer 13241 (CAS, CR, DUKE). Limon: Canton de Talamanca,
Bratsi; Alto Lari, Entre Surayo y Dapari, 50 m N de la desembocadura del Rio Dapari (Pare), junto al
Rio Lari, 9°25'10”N, 83°03'00”W, 300 m, 25 Feb. 1992, Aguilar & Schmidt 961 (INB, MO); Suerre,
Santa Clara, 300 m, Feb. 1896 (no exact day), Donnell-Smith 6554 (F, US); Zona Protectora Barbilla
on W side of plateau separating headwaters of N fork of Rio Danta from headwaters of Quebrada
Barreal, Rio Barbilla drainage (SE of Siquirres), 10°05'N, 83°28.5'W, 600-660 m, 11 Jan. 1987,
Grayum et al. 7932 (CAS, CR, MO); Canton de Talamanca, Bratsi, Amubri, Alto Lari, Kivut.
Quebrada innominada, margen derecha del Rio Dapari 9°24'20”N, 83°05'35”W, 1000 m, 11 Mar.
1992, Herrera 5294 (CAS, CR, INB, MO).

DISCUSSION. — Topobea dimorphophylla has leaf blades that are markedly unequal in size at
each node and copiously pubescent on both surfaces (Fig. 2A). It is most like 7. intricata which has
leaf blades that are only somewhat unequal in size at each node, longer floral peduncles (3.5—5.2 cm
vs. 0.8—1.4 in T. dimorphophylla), and free (vs. basally fused for 3.5—5 mm) floral bracts. In habit,
indument details, and the pronounced foliar dimorphism at each node, 7. dimorphophylla is also simi-
lar to T. tetramera which is readily separated by its 4-merous flowers, 4-locular ovary, completely
free inner and outer floral bracts, and truncate anther pores.

One collection of 7. dimorphophylla, Donnell-Smith 6554, was erroneously cited as a representa-
tive specimen of Clidemia costaricensis Cogn. & Gleason ex Gleason (Gleason 1939:126). This ap-
pears to have been an inadvertent error because this specimen, which is not mixed with C.
costaricensis, has young fruiting hypanthia with attached floral bracts that Gleason surely would have
recognized as something other than Clidemia.

ETYMOLOGY. — The epithet dimorphophylla is derived from the Latin word dimorphus, having
two forms, and the Greek word phyllus, relating to leaves, in reference to the pronounced difference in
leaf size at each node.

526 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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8. Topobea dodsonorum Wurdack, Phytologia 38:304. 1978.

TYPE. — ECUADOR. Los Rios/Pichincha border: near La Centinela at Km 12 on road from Pa-
tricia Pilar to Flor de Mayo, Montana de Ila, 600 m, 16 Jul—11 Aug. 1977, Dodson & Dodson 6752
(holotype: US!; isotype: MO, SEL).

Viny epiphytic shrub often with pendent branches 1—1.5 m long. Uppermost branchlets terete
with elevated interpetiolar ridges. Upper internodes, juvenile leaves, peduncles, bracts, and calyx
lobes caducously scurfy-puberulent with branlike or stellulate hairs. Mature leaves of a pair markedly
dimorphic in size; blades subcoriaceous, the larger at each node 7.4—19.5 = 3.6—9.5 cm, elliptic to
ovate or subcordate, the apex acuminate, the base broadly rounded and subpeltate, the margin entire,
5—7-nerved with well-developed pocket domatia formed at the point where the innermost primaries
diverge from the median vein, the transverse secondary veins spaced 1-2 mm apart at the widest por-
tion of the blade on the abaxial surface; petiole 3—5 mm long; the smaller blade 1.4—5 x 0.8-2.8 cm,
narrowly elliptic, the apex acuminate, the base rounded, the margin entire, 3—S-nerved with domatia
like those of the larger leaves but smaller in all dimensions and less conspicuous, the transverse sec-
ondary veins not conspicuous on the abaxial surface; petioles 0.1—1.3 mm long. Flowers erect, soli-
tary or borne in clusters of 2 to 3 in each leaf axil of distal branches; peduncles 1—2.5 cm long. Floral
bracts fused basally for 0.5—1 mm and much shorter than the hypanthium and calyx; outer bracts
22.5 x 2mm, ovate; inner bracts 2 x 2 mm, oblate to semicircular. Hypanthium (at anthesis)
campanulate, 4 x 3—4 mm. Calyx tube I—1.5 mm long; calyx lobes 1—1.8 x 2 mm, triangular. Petals 6,
glabrous, 5—6 x 2—3.5 mm, yellow, yellow-brown or greenish brown, oblong-obovate to subspatulate
with a blunt acute apex. Stamens 12; filaments 2—3.5 mm long, declinate and glabrous; anthers free,
1.75—2 x 0.5 mm, yellow, linear-oblong to somewhat subulate, each opening by a solitary, dor-
sally-inclined apical pore; connective thickened dorsally and prolonged dorso-basally into a blunt ap-
pendage 0.25 mm long. Ovary 1/2-inferior, 4-locular, somewhat elevated and rounded up to the stylar
scar but lacking a cone and collar. Style glabrous, 7 mm long; stigma punctiform. Berry 5—7 x
6—7 mm, greenish flushed with maroon at maturity. Seeds cuneate to narrowly pyriform, | mm long,
beige with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Often locally common in rainforests and low elevation
cloud forests of western and north-central Panama and eastern Ecuador at 540—1200 m. Flowering
collections have been made in January, February, April, June, July and September, fruiting during
these same months and in November.

REPRESENTIVE SPECIMENS EXAMINED. — PANAMA. Chiriqui: Edwin Fabrega Dam and Re-
serve in Fortuna along trail to hydrological station along Rio Hornito, 8°45’N, 82°5’W, 20 Jan. 1989,
Almeda et al. 6351 (CAS, PMA). Comarca de San Blas: Cerro Brewster, 9°18’N, 79° 16'W, 21 Apr.
1985, de Nevers et al. 5459 (CAS, MO, PMA).

DISCUSSION. — The diagnostic features of 7. dodsonorum include its pronounced foliar size di-
morphism at each node, well-developed foliar pocket domatia, triangular calyx lobes, and 4-locular
ovary. In his account of this species for Flora of Ecuador, Wurdack (1980) stated that domatia are
lacking in the smaller leaf at each node. Although the domatia on the smaller leaf are smaller in all di-
mensions, I have found them to be present on all specimens examined from throughout the range of
the species.

9. Topobea fragrantissima Almeda, Proc. Calif. Acad. Sci. 46:318. 1990.
TYPE. — PANAMA. Chiriqui: vicinity of Fortuna Dam, along trail across valley of Rio Hornito,

elev. 1100-1250 m, 12 Mar. 1988, Almeda et al. 6086 (holotype: CAS!; isotypes: CR!, F!, MO!,
PMA!, TEX!, US!).

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 527

Epiphytic or terrestrial shrubs or small trees | .5—4 m tall. Distal branchlets subquadrate and gla-
brous with interpetiolar ridges or lines. Vegetative buds and young leaves sparingly and caducously
lepidote-furfuraceous. Mature leaves of a pair equal to somewhat unequal in size, glabrous through-
out, 1.5—-5.5 cm long and 1.6—3 cm wide, elliptic to elliptic-obovate, the apex acuminate, the base
acute, the margin entire, 3-nerved or 3-plinerved abaxially with an additional submarginal pair of in-
conspicuous veins, secondary veins spaced mostly 0.25 mm apart at the widest portion of the blade.
Flowers erect, solitary or paired in leaf axils of upper branches; peduncles 2—3 cm long, glabrous;
outer bracts 5—11 x 3-5 mm, free, elliptic or rarely varying to obovate, glabrous, apex rounded; inner
bracts 4—6 x 4-5 mm, free, obovate, glabrous, apex broadly rounded. Calyx tube | mm long; calyx
lobes | mm long and !—1.5 mm wide basally, ovate to deltoid-ovate with a blunt callose-thickened
tooth on the abaxial apex of each lobe, margin entire but sometimes roughened along interlobe si-
nuses, glabrous on both surfaces. Petals 6, glabrous, |.2—1.4 x 1—1.2 cm, white flushed with pink uni-
laterally, obovate, apically rounded, entire. Stamens 12, free and declined to one side of the flower
opposing the style; filaments 5-6 mm long; anthers 3.5-5 mm long, | mm wide, yellow, lin-
ear-oblong and tipped with a solitary, dorsally-inclined pore; connective modified dorso-basally into
a deflexed spur 0.25 mm long. Ovary 1/2-inferior, 4-locular, glabrous at the apex but not elevated into
a cone or collar. Style declinate and sigmoid, glabrous, 10-11 mm long; stigma punctiform. Berry
7-10 x 10 mm. Seeds clavate to narrowly pyriform or pyramidate, | mm long.

DISTRIBUTION AND PHENOLOGY. — Local and uncommon in cloud forests from the Boquete re-
gion of W Panama (Chiriqui) to the Fortuna Dam region and E to Cerro Colorado at 1000-1300 m.
Flowering and fruiting material has been collected in January, March, April, and July.

REPRESENTATIVE SPECIMENS EXAMINED. — Chiriqui: Edwin Fabrega Dam and Reserve along
trail to Rio Hornito above Los Planes, 8°45'N, 82° 15’W, 18 Jan. 1989, Almeda et al. 6309 (CAS, MO,
PMA); Monte Rey, above Boquete, 21 Jul. 1971, Croat & Porter 15692 (CAS, MO). Chiriqui/Bocas
del Toro border; windswept cloud forest off the road to Cerro Colorado, 26 Jan. 1989, Almeda et al.
6418 (CAS, CR, DUKE, MO, PMA, US).

DISCUSSION. — The outstanding features of 7. fragrantissima are its completely glabrous leaves
with closely spaced (0.25 mm) transverse secondary veins on abaxial foliar surfaces, linear-oblong
anthers with solitary pores, and 4-locular ovary. Live flowers of this species produce a pleasant per-
fume-like fragrance, but nothing 1s known about tts role in attracting pollinators. For illustrations of
this species see Almeda (1990:319) and Plate 1.

10. Topobea gerardoana Almeda, sp. nov. (Fig. 3)

TYPE. — COSTA RICA. Limon. Cordillera de Talamanca between Quebrada Kuisa and Rio
Lari, 09°20'25”N, 83°13'45”W, elev. 2100 m, 17 Mar. 1993, Herrera 5914 (holotype: CAS!:
isotypes: CR, INB!, MO).

Frutex epiphyticus. Petioli 1.5—3.6 cm longi; lamina 6—12.5 x 2.1—5.5 cm elliptico-obovata vel
elliptica apice acuminata basi acuta, supra glabra, subtus sicut ramuli sparsiuscule pilis induti,
3—5-plinervata, nervis secundariis 1-3 mm inter se distantibus. Flores 6-meri in quoque nodo
superiore singuli vel bini, pedunculis 1.3—2 cm longis; bracteae exteriores 7-8 x 5—6 mm ovatae vel
elliptico-ovatae ca. 2-3 mm coalitae apice acuto vel rotundato; bracteae interiores 6-7 <x 6mm
omnino liberae. Calycis tubus 0.5—0.75 mm longus, lobis 5 mm longis. Petala alba, 1—1.2 x 0.8—1 cm
obovata apice mucronato. Filamenta 3—3.5 mm longa; antherae 3 x | mm inter se non cohaerentes,
poro unico dorsaliter inclinato; connectivum ad basim dorsaliter dente 0.25 mm longo descendenti
armatum. Stylus 1—1.1 cm; ovarium 6-loculare et 2/3 inferum apice glabro (cono ca. | mm alto).

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I Shwa_ i9qs

FiGurE 3. Topobea gerardoana Almeda. A. habit, ca. 1/3; B. representative leaf (abaxial surface), 3/5; C. enlarged leafbase
(abaxial surface) showing indument detail, ca. 1; D. young fruiting hypanthium with decussate floral bracts, 3; E. petal
(adaxial surface), 4; F. stamens, dorsal view (left) and profile view (right), ca. 7. (A-F from Herrera 5914.)

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 529

Epiphytic shrub. Uppermost internodes, vegetative buds, and abaxial surfaces of some mature
leaves sparingly covered with a caducous indument of rusty brown subulate and conic hairs. Mature
leaves of a pair mostly unequal in size; petioles 1.5—3.6 cm long; mature blades subcoriaceous,
6—12.5 cm long and 2.1—5.5 cm wide, elliptic-obovate to elliptic, the apex acuminate, the base acute,
the margin entire to obscurely denticulate, 3—S-plinerved with the innermost pair of primary veins di-
verging from the median vein 1-5 mm above the blade base, the subparallel secondary veins spaced
1—3 mm apart at the widest portion of the blade. Flowers erect, solitary or paired in leaf axils of upper-
most branches; peduncles | .3—2 cm long, subquadrate and lenticellate. Floral bracts entire, glabrate or
sparingly covered abaxially with a lanate indument, the margins ciliate or fimbriate; outer bracts 7-8
x 5—6 mm, fused at the base for 2-3 mm, concave, ovate to elliptic-ovate, the apex bluntly acute to +
rounded; inner bracts 6—7 x 6 mm, free, oblong to oblong-ovate, the apex rounded. Hypanthium (in
young fruit) suburceolate. Calyx tube 0.5—0.75 mm long; calyx lobes (in young fruit) erect, 5 mm
long and 3 mm wide at the base, triangular, irregularly and sparingly covered with a caducous lanate
indument on both surfaces. Petals 6, 1—1.2 x 0.8—1 cm, white, obovate, apex mucronate, entire. Sta-
mens 12; filaments 3—3.5 x 1.5 mm, complanate, glabrous and tapering to the apex; anthers 3 mm
long and | mm wide, free, yellow, oblong and somewhat sickle-shaped in profile view with a solitary
dorsally declined apical pore; connective thickened dorsally and modified dorso-basally into an ap-
pendage 0.25 mm long or less. Ovary 2/3-inferior, 6-locular, apex glabrous and elevated into a blunt
cone about | mm high. Style straight, glabrous, |—1.1 cm long; stigma punctiform. Immature berry 8
x 7-8 mm. Seeds narrowly pyriform, | mm long, tan with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Presently known only from cloud forests in southeastern
Costa Rica in a region upslope and south of the Valle de Talamanca at 1900-2100 m. The two known
collections were made in March. One is in flower, the other is in young fruit.

PARATYPE. — COSTA RICA. Limon: Canton de Talamanca. Bratsi, Amubri, Alto Lari, Kivut.
Ridge between Rio Lari and Rio Dapari, 9°22'45”N, 83°06'1S”W, 25 Mar. 1992, Herrera 5492
(CAS, INB).

DISCUSSION. — This little-known Costa Rican endemic has well-defined triangular calyx lobes
that are irregularly and sparingly covered with a caducous lanate indument (Fig. 3D), free anther
thecae that are somewhat sickle-shaped in profile view with a solitary apical pore (Fig. 3F), and an an-
ther connective that is modified dorso-basally into an appendage 0.25 mm long or less. Topobea
gerardoana does not appear to be particularly close to any congener with single-pored anthers.

ETYMOLOGY. — This species is named for Gerardo Herrera, stellar collector of Costa Rican
plants.

11. Topobea intricata Almeda, Brittonia 53:157. 2001.

TYPE: COSTA RICA. Cartago: Highway #224 on property of ICE hydroelectric plant (now
Tapanti National Park) ca. 20-24 km E of the church in Orosi, elev. 1500-1800 m, 5 Jan. 1974, F.
Almeda et al. 2366 (holotype: CAS!; isotypes: BM!, CR!, DUKE!, INB!, MEXU!, MO!, NY!).

Epiphytic or terrestrial shrub 1—2 m tall with spreading branches to 3 m long. Uppermost branch-
lets rounded. Cauline internodes, vegetative buds, leaf blades (both surfaces), peduncles, floral bracts,
hypanthia, and calyx lobes moderately to copiously hirsute with smooth ferrugineous hairs mostly
0.5—3(-6) mm long. Mature leaves of a pair typically unequal in size; blades thin and papery when
dry, the larger one at each node 7.5—17.2 x 3.5—7.5 cm, the smaller one 1.8—10 x 1.7—5.4 cm, elliptic
to elliptic-ovate, the apex acuminate to caudate-acuminate, the base obliquely obtuse to rounded, the
margin denticulate to subentire (sometimes obscurely and remotely so), 3—5-plinerved with the inner-
most pair of primary veins diverging from the median vein |—5 mm above the blade base, the trans-
verse secondary veins spaced 2—6 mm apart at the widest portion of the blade. Flowers erect or

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horizontally spreading, solitary in the axils of uppermost branches; peduncles 3.5—5.2 cm long, terete.
Floral bracts green and entire; outer bracts 2.5—3.2 x 1.7-2.6 cm, free, ovate to subcordate, the apex
acute to acuminate, trinerved; inner bracts 1.9—2.5 x 0.7—1 cm, the basal half closely appressed to but
free from the hypanthium, narrowly elliptic-lanceolate, the apex acute to acuminate. Calyx lobes (at
anthesis) linear oblong, 6-7 mm long and | mm wide at the + deltoid base between sinuses. Petals 6,
glabrous, 1.6—2.3 x 1.4-1.7 cm, white flushed with pink distally on the abaxial surface, thin and trans-
lucent when dry, obovate, apex rounded, base somewhat clawed, entire. Stamens 12; filaments 5—6.5
x | mm, declinate, complanate, glabrous; anthers free, 5—7.5 x | mm, yellow, oblong-subulate and
incurved distally, each with 2 confluent, dorsally-inclined pores at the truncate apex; connective
thickened dorso-basally into a blunt deflexed appendage 0.5—1 mm long. Ovary 1/2-inferior (at
anthesis), 6-locular, glabrous at the summit which is elevated into a lobulate collar 1—-1.5 mm high
surrounding the style base. Style glabrous, 0.9—-1.4 cm long; stigma punctiform. Berry red at maturity,
1-1.7x 1.3-1.4 cm. Seeds mostly | mm long, beige, cuneate to narrowly pyriform.

DISTRIBUTION AND PHENOLOGY. — Locally common in cloud forests of Tapanti National Park
in central Costa Rica and the Fortuna region of western Panama at 1 100—1800 m. Collected in flower
from December through June, in fruit from March through July.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Cartago: About 15km S of
Tapanti along new road on the E slope above the Rio Grande de Orosi near the concrete bridge,
9°42'N, 83°47'W, 12-17 Dec. 1969, Burger & Liesner 6810 (CAS, CR, F); Canton de Paraiso,
Parque Nacional Tapanti, Valle de Reventazon, Sendero Arboles Caidos, 09°45'00”N, 83°47'00"W,
21 July 1994, Quesada 123 (INB). PANAMA. Bocas del Toro: Fortuna Dam area, along continental
divide trail bordering Chiriqui Province, 08°45'04”N, 82°15'04’W, 10 Mar. 1988, Almeda et al.
6048 (BM, CAS, CR, MEXU, MO, NY, PMA); 2 km W of continental divide along trail to elfin for-
est, 08°47'N, 82°13'W, 26 Mar. 1985, Hampshire & Whitefoord 981 (BM, CAS). Chiriqui: Fortuna
Dam area, trail to meteorological station of Rio Hornito, 08°45'N, 82°18'W, 23 June 1994, Croat &
Zhu 76310 (CAS, MO); Fortuna Dam area, road from Gualaca to Chiriqui Grande on continental di-
vide trail W of road, 08°45'N, 82°15'W, 18 Jan. 1986, de Nevers & McPherson 6852 (CAS, MO);
Road from Fortuna Lake to Chiriqui Grande on trail W of continental divide, 08°47'N, 82°13'W, 22
Mar. 1985, Hampshire & Whitefoord 850 (BM); Cordillera Central, 7 Dec. 1996, Montenegro 1585
(CAS, SCZ); Distrito Boquete, Fortuna Dam site along trail following continental divide, 8 Feb. 1985,
van der Werff & van Hardeveld 6707 (CAS, MO).

DISCUSSION. — Topobea intricata has a copious cover of ferrugineous hairs throughout, thin pa-
pery leaves, solitary floral peduncles, and ovate to subcordate outer floral bracts. It most closely re-
sembles 7. dimorphophylla in habit, pubescence details, leaf morphology, and petal color. For a
comparison of these two species see the discussion under the latter. Topobea intricata is also superfi-
cially similar to Blakea wilburiana Almeda of central Panama. The latter has elliptic-lanceolate outer
floral bracts, filiform to nearly acicular calyx lobes, and smaller anthers (4—5 x 1—2 mm) that are ob-
long and laterally compressed with ventrally-inclined (vs. dorsally-inclined) apical pores.

As noted in the protologue, what may be a regional variant or a closely related undescribed taxon,
represented by Herrera 6036 (CAS, INB) and Herrera 2832 (CAS), has been collected on the Carib-
bean slopes of the Cordillera de Talamanca in southern Costa Rica. This entity, which is known from
five gatherings, is similar to T. intricata in all details but has outer floral bracts that are uniformly ellip-
tic and consistently smaller (1.3—1.5 x 0.8—1.5 cm). The known collections of T. intricata from geo-
graphically distant areas in Costa Rica and Panama are homogeneous morphologically. It is for this
reason that I am reluctant to consider this distinctive population a mere variant without additional
study. See Almeda (2001:158) for an illustration of this species.

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 531

12. Topobea laevigata (D. Don) Naudin, Ann. Sci. Nat. Bot. 3, 18:150. 1852.

Blakea laevigata D. Don, Mem. Wern. Nat. Hist. Soc. 4:327. 1823. TYPE. — MEXICO, without exact locality,
Sessé & Mocino s.n. (BM! ex Herb. Lambert).

Epiphytic shrub or tree S—10 m tall with a lateral spread of up to 8 m. Uppermost branchlets
rounded-quadrate becoming rounded with age, essentially glabrous throughout, the young floral buds
and peduncles sometimes covered with a caducous furfuraceous indument of stellulate or amorphous
branlike hairs. Mature leaves of a pair equal to slightly unequal in size; petioles 0.8—3.4 cm long;
blades coriaceous, 3.6—18 x 1.8—7.3 cm, elliptic to elliptic-obovate, the apex acuminate to abruptly
short-acuminate, the base acute to obtuse, the margin entire, 3—5-nerved with the marginal pair of
veins obscure or lacking or 3—5-plinerved with the inner pair of primary veins diverging from the me-
dian vein 4—7 mm above the blade base (on abaxial surface), commonly with some leaves bearing
slitlike pit domatia at the abaxial margin of the blade near the petiole-laminar junction, the transverse
secondary veins often obscure or 0.5—1 mm apart at the widest portion of the blade when visible.
Flowers erect, solitary or in clusters of 24 in each leaf axil of distal branches; peduncles 1.2—2 cm
long. Floral bracts glabrous at maturity, + rigid, concave, and tightly enveloping the hypanthium,
outer bracts 4.5—-5 x 6-7 mm, fused basally for 2-4 mm or sometimes essentially free to the base,
broadly deltoid to oblate or semicircular; inner bracts 4 5—6 mm, free, semicircular. Hypanthium (at
anthesis) campanulate, 5-6 x 6-7 mm. Calyx tube 2.5 mm long, erect and cupulate, calyx broadly
flattened into low undulations 0.5 x 3 mm. Petals 6, glabrous, 1.3 x 0.4-0.5 cm, white, narrowly
obovate. Stamens 12; filaments 4-5.5 mm long, declinate, complanate, glabrous; anthers free, 4.5—5
x 1 mm, yellow, each with 2 confluent, dorsally-inclined pores at the emarginate apex; connective
thickened dorsally, unappendaged or with a blunt dorso-basal callosity. Ovary 1/2-inferior, 6-locular,
elevated apically into a low flangelike rim that forms a wide circle around the stylar scar. Style gla-
brous, 1.2—1.3 cm long; stigma punctiform. Berry 6—7 x 6 mm. Seeds ovoid to + pyriform, | mm long,
beige with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Local in montane rainforests, seasonal evergreen forests
and oak-pine forests from southern Mexico (Pueblo and Veracruz, southward) to Guatemala and
Belize at 300—1850 m. Flowering collections have been made in January, April, July, September, Oc-
tober, and December, fruiting collections have been gathered in every month except July and Septem-
ber.

REPRESENTATIVE SPECIMENS EXAMINED. — BELIZE. Toledo: high ridge, Jacinto Creek, Rio
Grande, 28 Oct 1944, Gentle 4928 (CAS, LL). GUATEMALA. Alta Verapaz: Sebol, in potrero, 21
Apr. 1964, Contreras 4440 (DS). Izabal: El Estor, in high forest, Contreras 11464 (CAS). Peten: La
Cumbre, top of hill in zapotal, on Pusila Road, 5 km north, Lundell & Contreras 20228 (CAS, LL).
MEXICO. Chiapas: Municipio of Berriozabal, 13 km N of Berriozabal near Pozo Turipache and
Finca El Suspiro, 25 Dec. 1972, Breedlove & Thorne 30771 (DS): Municipio of Cintalapa, between
Colonia Francisco I. Madero and Colonia A. Lopez Mateos, 29 Mar. 1981, Breedlove 50564 (CAS):
Municipio of La Trinitaria, 10 km ENE of Dos Lagos above Santa Elena, 19 Jan. 1982, Breedlove &
Almeda 57558 (CAS); Municipio of Ocosingo, adjacent to Laguna Ocotal Grande, 6 Feb. 1973,
Breedlove 33070 (DS): Municipio of Peltalcingo, steep slope of Ahk’ulbal Nab above Peltalcingo, 28
Mar. 1981, Breedlove 50424 (CAS). Oaxaca: Municipio San Miguel Chimalapa. Cerro Salomon ca.
2 km en linea recta al NNO del Cerro Guayahitos, ca. 43 km en linea recta al N de San Pedro
Tapantepec, 23 Dec. 1985, Wendt et al. 5156 (CAS). Puebla: Municipio Eloxochitlan, adelante de San
Miguel Eloxochitlan en camino nacia El Mirador, 12 June 1985 Chazaro & Leach 3392 (CAS).
Veracruz: Municipio Hidalgotitlan. A fluente del Rio Las Cuevas, + 5 horas a pie al S de la Laguna, 16
Apr. 1982, Wendt et al. 3845 (CAS).

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DISCUSSION. — When present in a species of Topobea, acarodomatia are commonly produced
with consistency on most leaves of all individual plants. In 7. /aevigata, the production of domatia is
inconsistent. In the specimens examined for this study, no foliar domatia were found in collections
from Belize and Guatemala. Among Mexican collections studied, domatia were observed as follows:
Chiapas—some leaves on 18 of 20 collections; Oaxaca—some leaves on two of five collections;
Puebla—some leaves on the single collection studied; Veracruz—some leaves on three of four collec-
tions examined. For differences between 7. /aevigata and T. calyvcularis see the discussion under the
latter.

13. Topobea lentii Almeda, Brittonia 53:160. 2001.

TYPE: COSTA RICA. Cartago: 3 km E of Cachi, beside Rio Naranjo, 1300 m, 11 July 1971, Lent
2000 (holotype: MO!; isotypes: BM!, CR!, F!, DUKE!, PMA!, US!).

Epiphytic shrub. Uppermost branchlets bluntly quadrate with elevated internodal lines or ridges
but becoming rounded and coarsely striate with age. Young vegetative buds, floral peduncles, young
floral bracts, hypanthia, and calyx lobes moderately covered with a caducous indument of scurfy
and/or stellulate hairs. Mature leaves of a pair equal to somewhat unequal in size, essentially glabrous
on both surfaces; blades coriaceous, 4.2—9.1 cm long and 1.94.3 cm wide, elliptic to elliptic-ovate,
the apex bluntly acuminate, the base acute to obtuse, the margin entire, 5-plinerved, the innermost pri-
maries diverging from the median vein 2-6 mm above the blade base (on abaxial surface) and forming
poculate acarodomatia sparsely to moderately covered with barbellate hairs in the angles formed with
the median vein, the transverse secondary veins spaced 0.25 mm apart at the widest portion of the
blade. Flowers erect, 2—5 per axil in uppermost leafy branches; peduncles 5—7 mm long, lenticellate.
Floral bracts free, sometimes recurved at the apex; outer bracts 3—3.5 x 22.5 mm, elliptic-ovate, the
apex bluntly acute to rounded; inner bracts 2—2.5 x 2—2.5 mm, elliptic to elliptic-ovate, the apex
rounded. Calyx tube 1.5—2 mm long, cupulate; free portions of calyx lobes 0.5—1.5 mm long and
0.5—1.5 mm wide basally between sinuses, broadly subtruncate and often apiculate, somewhat
callose-thickened at the median apex abaxially. Petals 6, glabrous but sometimes fringed with incon-
spicuous caducous white hairs, 6—6.5 x 2-3 mm, reportedly white (fide Lent 2000) or pink (fide
Haber ex Bello 5174), obovate-rhombic, the apex bluntly long-acuminate and somewhat concave.
Stamens 12; filaments 3 mm long, somewhat declinate, complanate, glabrous; anthers free or laterally
connate (in part), 2.5 mm long, 0.25 mm wide, pale yellow (?) and granulose along the lower ventral
half of the thecae, subulate, each with a solitary, dorsally-inclined apical pore; connective thickened
dorso-basally and prolonged near the filament insertion into a deflexed caudiform appendage 0.5 mm
long. Ovary 1/2 inferior, 6-locular, glabrous and elevated at the apex into a lobulate collar 1 mm high
that surrounds the style base. Style glabrous, 5-7 mm long; stigma punctiform. Berry 3-7 x
2.5—7 mm. Seeds 0.75—1 mm long, beige, cuneate to obovoid or narrowly pyriform.

DISTRIBUTION AND PHENOLOGY. — A rare cloud forest epiphyte known only from the vicinity of
Cachi in central Costa Rica and the Rio Chiquito de Tilaran in Guanacaste province at 1300-1450 m.
Collected in flower in July and December, in fruit in July.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Cartago: pasture on hilltop, 2 km E
of Cachi, 9°49’N, 83°47'W, 16 Dec. 1972, Lent 3118 (F, US). Guanacaste: Rio Chiquito de Tilaran,
Rio Negro, 10°22'N, 84°52'W, 1 July 1986, Haber ex Bello 5174 (CAS).

DISCUSSION. — Notable features of 7. Jentii are its hair tuft foliar domatia, small fruiting
hypanthia, clawed obovate-rhombic petals, ventro-basally granulose anther thecae, and dorso-basal
caudiform appendages on anther connectives. It most closely resembles the widespread 7. watsonii
which differs in its well-defined triangular calyx lobes, larger petals (10—16 x 4.5—7 mm), longer an-

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 533

ther thecae (6—7.5 mm) that are connate for a major portion of their length, and the absence of foliar
domatia. See Almeda (2001:161) for an illustration of this species.

14. Topobea maurofernandeziana Cogn., DC. Monogr. Phan. 7:1193. 1891.

TYPE. — COSTA RICA. Foréts de Juan Vifias, 25 Jan. 1890, Tonduz 1844 (holotype: BR!:
isotype: CR!).

Topobea durandiana Cogn., Bull. Soc. Roy. Bot. Belgique Ser. 3, 30:268. 1892. TYPE. — COSTA RICA. Bord
d’un torrent a Buenos Aires, 250 m, II. 1891, Pittier 3789 (holotype: BR!).

Blakea intercepta Gleason, in Woodson & Schery, Ann. Missouri Bot. Gard. 28:435. 1941. TYPE. — COSTA
RICA, without exact locality, 20 June 1874, O. Kuntze s.n. (holotype: NY!).

Terrestrial or epiphytic shrub 2—5 m tall and 3—7 m in diameter. Uppermost cauline internodes
quadrate. Uppermost internodes, young vegetative and floral buds, and petioles sparsely to densely
furfuraceous pubescent but glabrate with age. Leaves thick and coriaceous when dry; petioles
1.8—9 cm long; blades 11.5—23 x 7—18.5 cm, elliptic to elliptic-oblong or ovate, the apex abruptly
acuminate, the base obtuse to rounded, the margin entire, 5-nerved or 5-plinerved, adaxially glabrous,
sparingly furfuraceous to almost glabrous abaxially. Flowers 2—S (rarely solitary) in each leaf axil of
upper branchlets; peduncles 0.6—2.5 cm long. Inner and outer floral bracts 0.7—1.2 x 0.7—1.5 cm, es-
sentially free, ovate to suborbicular, the apex rounded to retuse or obscurely apiculate. Calyx tube (on
fruiting hypanthia) 0.6—0.8 cm long from the torus; free portions of calyx lobes | x 5 mm, broadly de-
pressed-truncate with an apiculate callosity at the median apex, margin + entire. Petals 6, glabrous,
1.7—2.5 x 1—1.3 cm, pink, obovate, the apex rounded to obliquely subtruncate. Stamens 12; filaments
7-9 mm long, declinate; anthers laterally connate, 6—9 x 1-2 mm, pink or yellow flushed with pink or
red distally, linear-oblong and incurved distally, each with 2 confluent, dorsally-inclined apical pores;
connective prolonged dorso-basally into a caudiform appendage | mm long. Ovary 1/3 inferior,
6-locular, the apex covered with a caducous ring of minute glandular hairs surrounding the stylar scar.
Style glabrous, 1.2—1.5 cm long; stigma truncate to capitellate. Berry 1—-1.4 x 1.3—1.5 cm. Seeds
1 mm long, beige, cuneate to clavate.

DISTRIBUTION AND PHENOLOGY. — Locally common in rainforests and cloud forests, often on
remnant pasture trees from southern Mexico (Guerrero and Oaxaca) disjunct to Nicaragua, Costa
Rica, and Panama from sea level to 1600 m. Flowering collections have been made from September
through July, fruiting collections from February through November.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela: about 2-4 km N of
Bijagua on road to Upala, 1 Mar. 1978, Almeda & Nakai 4041 (CAS, CR). Cartago: about 3 km E of
Cachi, 24 Jul. 1977, Almeda et al. 3209 (CAS, CR). Guanacaste: Parque Nacional Guanacaste,
Estacion Cacao, Liberia, 10°55'45”N, 85°28'15”W, 2 Nov. 1990, Chavez 339 (CAS, CR, INB, MQ).
Heredia: Parque Nacional Braulio Carrillo, Estacion Magsasay. Sarapiqui, 10°24'10°N,
84°03'30"W, 4 Apr. 1991, Aguilar 126 (CAS, CR, INB, MO). Limon: Parque Nacional Tortuguero,
Estacion Cuatro Esquinas. Laguna Tortuguero, 10°32'02”N, 83°30'26”W, 4 Jul. 1990, Chavarria
116 (CAS, CR, INB, MO). Puntarenas: Las Cruces Tropical Botanical Garden and vicinity about
6.4 km S of San Vito de Java, 18 Mar. 1978, Almeda et al. 4273 (CAS, CR). San José: Reserva
Bioldgica Carara. Sitio El Chuzazo, 9°45'05”N, 84°31'50”°W, 14 Feb. 1990, Zuniga 110 (CAS, CR,
INB, MO). MEXICO. Guerrero: Sierra Madre Sur. Along road between El Paraiso and Puerto del
Gallo, 6—8.7 miles NE of El Paraiso, 9 Mar. 1987, Daniel & Bartholomew 4932 (CAS). Oaxaca:
Distrito de Putla, La Cascada, 5 kmal NE de La Hacienda, sobre al camino San Vicente-San Isidro, 13
Apr. 1987, Garcia et al. 3171 (CAS). NICARAGUA. Chontales: vicinity of Finca San Pedro de
Oluma, on NE flanks of Cerro Oluma, 12°18'N, 85°23'W, 22 Sep. 1983, Nee 28328 (CAS).

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Matagalpa: falda norte del Cerro Musun, frente a trocha a Wanawas, 16 May 1980, Araquistain &
Moreno 2798 (CAS). Rio San Juan: near Canto Chontalefio, 20 km NE of El Castillo, 18-21 Apr.
1978, Neill & Vincelli 3560 (CAS). Zelaya: NW de Estacion Experimental El Recreo, 12°10'N,
84°18'W, 2 May 1982, Sandino 2702 (CAS). PANAMA. Chiriqui: 15-20 km S of Volcan, road to
Barrilos, 22 Dec. 1974, Dressler 4903 (CAS).

DISCUSSION. — This is one of the most common species of Topobea in lowland sites of southern
Central America. Like 7. watsonii, the only notable variation in this species is in the amount and per-
sistence of the indument on upper internodes, vegetative and floral buds, petioles and peduncles. In
the protologue of 7. durandiana, which was published shortly after 7. maurofernandeziana,
Cogniaux gives no specific differences that can be used to separate these two species. Judging from
the species descriptions it appears that Cogniaux assigned those collections with
furfuraceous-puberulent petioles and peduncles to 7. durandiana whereas collections from the gla-
brous end of the spectrum were recognized as 7. maurofernandeziana.

Blakea intercepta is also included here in synonymy with confidence. The holotype consists of
three leaves and many young fruits. Although petals and anthers are lacking, the available material is
an exact match for 7. maurofernandeziana.

15. Topobea mcphersonii Almeda, Brittonia 53:163. 2001.

TYPE: PANAMA. Comarca de San Blas: San Blas boundary trail on Llano-Carti road, 9° 15'N,
79°00'W, elev. ca. 350 m, 27 Jan. 1986, McPherson & Merello 8176 (holotype: CAS!; isotypes: BM,
CR, EAP, MEXU, MO, PMA, US).

Epiphytic shrub. Uppermost branchlets bluntly quadrate becoming rounded with age. Young
vegetative buds covered with an amorphous lanate indument. Mature leaves of a pair equal to slightly
unequal in size, glabrous on both surfaces but glandular-punctate abaxially; petioles 0.5—1.2 cm long;
blades coriaceous, 4.1—7.5 cm long and 1.7—3 cm wide, elliptic-obovate to obovate, the apex bluntly
cuspidate, the base acuminate, the margin entire, 3—5-plinerved, the marginal pair of primaries incon-
spicuous and concealed by the revolute margins when dry, the transverse secondary veins spaced
0.5—1 mm apart at the widest portion of the blade. Flowers erect, solitary in leaf axils of uppermost
branches; peduncles 1.2—2 cm long. Floral bracts green flushed with red, entire and glabrous through-
out; outer bracts 1.6—2.1 x 1—-1.6 cm, free but conspicuously decurrent on and imparting a winged as-
pect to the the peduncle, elliptic to elliptic-obovate, apex bluntly cuspidate to bluntly acute; inner
bracts 1.1—1.5 x 0.3—-0.6 cm, free, linear-oblong to narrowly obovate and distinctly cucullate, apex
bluntly acute to obtuse. Calyx tube 2 mm long; free portions of calyx lobes 0.5—1 mm long and
34mm _ wide basally between sinuses, broadly depressed-triangular, margin entire,
callose-thickened at the median apex abaxially, glabrous throughout. Petals 6, glabrous, 2.1—2.7 x
1.2—-1.5 cm, reportedly white-pink, thin and translucent when dry, obovate, apically rounded, entire.
Stamens 12; isomorphic, filaments 5—6 mm long, declinate, complanate, glabrous; anthers laterally
connate for most of their length, 3-5 mm long, | mm wide, yellow, linear-oblong with each anther sac
opening by a separate, dorsally-inclined terminal pore; connective thickened dorsally and dilated
dorso-basally into an upturned or deflexed somewhat flattened, blunt appendage ca. 0.5 x 0.25 mm.
Ovary completely inferior, 6-locular, glabrous at the apex which is elevated into a blunt cone 1-3 mm
high. Style and stigma not seen. Berry subglobose, 0.6—0.7 x 0.7 cm. Seeds mostly | mm long, beige,
cuneate-clavate.

DISTRIBUTION AND PHENOLOGY. — A lowland rainforest species known only from El
Llano-Carti region of north-central Panama at 100-500 m. Collected in flower and fruit in January
and February. j

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 395

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Comarca de San Blas: Rio Nergala,
9°22'N, 79°7'W, 12 Jan. 1985, de Nevers & Herrera 4530 (CAS, MO). Panama: 14 km above
Panamerican Highway on road from El Llano to Carti-Tupile, 20 Feb. 1973 (fr), Kennedy 2502 (MO,
US).

DISCUSSION. — Topobea mcphersonii is unlike other described congeners in having floral bracts
that are prominently decurrent on the floral peduncles. The inner floral bracts, which are oblong and
do not tightly envelop the hypanthia, are also distinctive as are the laterally connate anther thecae with
cleft or divergent pores and upturned or deflexed staminal appendages. Because of these unique fea-
tures, the relationships of this species are obscure at this time. For an illustration of this species see
Almeda (2001:164).

16. Topobea multiflora (D. Don) Triana, Trans. Linn. Soc. London 28:149. 1871. (Fig. 4)

Blakea multiflora D. Don, Mem. Wern. Nat. Hist. Soc. 4:326. 1823. (Fig. 4) TYPE. — PERU. Without exact lo-
cality: Pavon 127-3 (holotype: K!, fragment at BR!).

Epiphytic shrub 2-4 m tall or free standing tree to 8 m tall. Uppermost branchlets quadrate be-
coming somewhat striate and rounded with age, the younger nodes covered with caudate-acuminate
stipuliform flaps ca. 3 mm long that envelop caducous tufts of hair, the distal portions of each flap +
caducous with age and leaving a prominent interpetiolar ridge or corky line. Vegetative and young
floral buds, calyx lobes (in bud), peduncles, and young pedicels moderately to sparsely covered witha
caducous indument of amorphous scurfy hairs. Mature leaves of a pair essentially equal in size, gla-
brous on both surfaces at maturity; petioles 1.4-6cm long; blades coriaceous, 10.3-18.5 =
6—10.4 cm, elliptic, the apex abruptly acuminate, the base acute, the margin entire, 5-nerved or
5-plinerved with the innermost primaries diverging from the median vein ca. 5 mm above the blade
base (on abaxial surface) with hair tuft acarodomatia that are sparsely to moderately covered with ro-
bust shaggy hairs (0.5—2 mm long) in the angles formed with the median vein, the transverse second-
ary veins spaced 0.25—1.5 mm apart at the widest portion of the blade. Flowers erect, borne in clusters
of (1—) 2-6 in each leaf axil of distal branches; peduncles 0.3—2.1 cm long. Floral bracts essentially
glabrous at maturity and closely enveloping the hypanthium; outer bracts S—8 x 5—7 mm, fused ba-
sally for 4-6 mm, broadly ovate to suborbicular, the free distal portion bluntly acute to broadly
rounded; inner bracts 7—9 x 10—11 mm, free but imbricate, obdeltoid to suborbicular, often appearing
+ rounded due to the firm concave posture. Hypanthium (at anthesis) campanulate, 7-8 x 5-6 mm.
Calyx tube 4-6 mm long, erect and cupulate; calyx subtruncate or barely evident as broadly oblate
apiculate lobes 0.5 4—5 mm. Petals 6, glabrous, |.1—1.8 x 0.5—1.2 cm, pink with a translucent whitish
inverted V-shaped lens at the base of each petal, narrowly obovate to spatulate, the apex bluntly acute
to + rounded, margin entire but inconspicuously caducous-fimbrillate. Stamens 12; filaments 9 mm
long, declinate, complanate, glabrous; anthers laterally connate for at least half their length, 6—9 =
1 mm, yellow, oblong-subulate, each with 2 confluent, dorsally-inclined apical pores; connective
thickened dorsally and prolonged dorso-basally into a deflexed toothlike appendage 0.5—2.5 mm
long. Ovary 1/4-inferior, 6-locular, elevated apically into a glabrous cone 3 mm high and a lobulate
collar ca. 1 mm long. Style glabrous, 1.3—1.5 cm long; stigma + truncate to somewhat expanded. Ma-
ture berry and seeds not seen.

DISTRIBUTION AND PHENOLOGY. — Locally common in rainforest habitats from central Costa
Rica to Panama, Ecuador, Peru, and Bolivia from sea level to 1400 m. Flowering has been recorded
from November through July; fruiting collections have been made in February and March and from
May through July.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela: Reserva Forestal de San
Ramon, 10°12'53”N, 84°36'28”W, 28 Jan. 1987, Herrera & Solis 457 (CAS, CR, MO). Puntarenas:

536 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

Sian

i ‘

F oes
yes
3 aS

Ly 5g

FIGURE 4. Topobea multiflora (D. Don) Triana. A. habit, 1/5; B. nodal enlargement showing stipuliform flaps. ca. 3; C. rep-
resentative leaf (abaxial surface), ca. %; D. enlarged leaf base (abaxial surface) showing hair tuft acarodomatia, ca. 3; E.
flower bud with enveloping bracts, ca. 3; F. petal (adaxial surface), ca. 3;G.androecium, ca. 3. (A, B from McPherson 7630;

C, D from McPherson 15046; E-G from Almeda et al. 4271 and McPherson 7964.)

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 537

Las Cruces Tropical Botanical Garden and vicinity about 6.4 km S of San Vito de Java, 18 Mar. 1978,
Almeda et al. 4271 (CAS, CR). San José: about 13-18 km S of San Isidro del General, 5 Mar. 1978,
Almeda & Nakai 4105 (CAS, CR, INB, MBM). PANAMA. Coclé: ca. 9 km beyond the market in El
Valle de Anton ona rock road to trail head to Cerro Caracoral, 16 Feb. 1996, Almeda et al. 7600 (CAS,
PMA); along Llano Grande to Coclesito road above Cascajal, near divide, 8°42’N, 80°28’W, 11 Jan.
1986, McPherson 7964 (CAS, MO, PMA); near El Valle de Anton, 8°37'N, 80°07'W, 25 Nov. 1985,
McPherson 7630(CAS, MO, PMA). Colon: Rio Guanche, |-4 km upstream from Portobelo Road, 10
Dec. 1973, Gentry 8767 (MO). Darién: Alturas de Nique on the Serrania de Pirre above the Cana min-
ing camp, | Mar. 1988, Almeda & McPherson 5967 (CAS, PMA); Cana region on trail above Cana
leading to ridge of Pirre massif, 7°45'N, 77°45'W, 5 May 1990, McPherson 15046 (CAS, MO,
PMA). Panama: road past Altos de Pacora, 3—3.5 miles NE of Altos de Pacora, 11.1—11.6 miles be-
yond Lago Cerro Azul, 9°15'N, 79°25'W, 19 June 1988, Croat 68669 (CAS, MO, PMA).

DISCUSSION. — This frequently collected species is recognized by a combination of characters
that includes stipuliform nodal flaps on uppermost branches (Fig. 4B), hair tuft domatia on abaxial fo-
liar surfaces (Fig. 4D), laterally connate anther thecae, and dorso-basal toothlike appendages on an-
ther connectives (Fig. 4G). In the past, Central American collections of 7. multiflora have been
erroneously identified as 7. calycularis. Standley (1938), for example, reported it from Costa Rica (as
T. calycularis) based on collections from Cafias Gordas made in 1897 (Pittier 10955 and Pittier
11062, both at US). It was not reported for Panama by Gleason (1958) because the species had not yet
been collected in that country. Topobea calycularis is presently known only from Mexico (Chiapas)
and adjacent Guatemala. Like 7. multiflora, it also has hair tuft foliar domatia but it has free anther
thecae that are essentially unappendaged, a 4-locular ovary, and lacks stipuliform nodal flaps.

For the most part, Central American material of 7. multiflora has modally shorter floral
peduncles (0.3—2.1 cm vs. 2—3 cm) than South American populations. Cogniaux was aware of this
difference and may have intended to give formal taxonomic recognition to the Central American pop-
ulations because he annotated at least one Costa Rican collection (Pittier 10955, BR) as T. multiflora
var. brevipedunculata. The fact that this infraspecific epithet was never published also suggests that
he may have changed his mind. I see no compelling reason for recognizing such an entity.

17. Topobea parasitica Aubl., Hist. Pl. Guiane Fr. 1:476. 1775.
TYPE. — FRENCH GUIANA. Sinnemary River, Aublet s.n. (holotype: BM!).

Topobea regeliana Cogn., DC. Monogr. Phan. 7:1085. 1891. TYPE. — PANAMA. Chagres, Isthmus of Pan-
ama, Mar. 1850, Fendler 295 (holotype: LE: isotypes, BM!, BR! [2 sheets], K!, MO!, US!).

Topobea praecox Gleason, Phytologia 3:355. 1950. TYPE. — PANAMA. Cocle: El Valle de Anton, vicinity of
La Mesa, Allen 2788 (holotype: NY!; isotypes: NY![2 sheets]).

Topobea membranacea Wurdack, Brittonia 9:108. 1957. TYPE.— COLOMBIA. Antioquia: Anori-Cruces
road, 11 May 1944, Core 677 (holotype: US!; isotype: NY!).

Robust shrub or small tree 2-10 m tall, often epiphytic or rupicolous. Uppermost branchlets
bluntly quadrate becoming rounded with age. Juvenile vegetative buds and uppermost nodes covered
with smooth stramineous hairs and a dense cover of stellulate or scurfy hairs. Uppermost cauline
internodes, petioles, peduncles, abaxial leaf surfaces (especially elevated primary veins), outer floral
bracts, and calyx lobes (in bud) moderately to sparsely covered with a varying mixture of scurfy,
stellulate, and roughened hairs that fall away to varying degrees with age. Mature leaves of a pair
somewhat unequal in size, glabrous on the adaxial surface at maturity, sparingly covered with scurfy
or stellulate hairs to nearly glabrous on the abaxial surface at maturity; petioles 1.8—9 cm long; blades
membranaceous to subcoriaceous, 6.2—25.5 x 5—16 cm, elliptic varying to elliptic-ovate, the apex

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abruptly short-acuminate, the base obtuse to broadly rounded but sometimes varying to acute, the
margin typically entire, rarely varying to denticulate, (S—)7(—9)-nerved with secondary veins spaced
1-4 mm apart at the widest portion of the blade, or S—7-plinerved in some plants and then the inner
pair of primary veins diverging 0.5—1 cm above the blade base (on abaxial surface). Flowers erect,
1-7 per leaf axil of uppermost branches; peduncles 0.3—1.9 cm long. Floral bracts free but appressed
to and commonly obscuring the hypanthium below the calyx; outer bracts 5-10 x 5—9 mm, broadly
ovate to subrotund, the apex typically rounded but varying to broadly obtuse, bluntly acute, acuminate
or somewhat emarginate; inner bracts 5—9 x 5—9 mm, ovate to subrotund, the apex typically broadly
rounded. Hypanthium (post anthesis) campanulate, 5—8 x 6-8 mm. Calyx tube 24 mm long, erect
and cupulate; free portions of calyx lobes 1|—2 long and 3-4 mm wide basally between interlobe si-
nuses, triangular varying to broadly depressed-triangular to nearly undulate in fruit. Petals 6, gla-
brous, 1.4—2 x 0.7—1.1 cm, reportedly pink, reddish-pink, or magenta, obovate, the apex rounded to
obtuse. Stamens 12; filaments 9-12 mm long, declinate, complanate, glabrous; anthers laterally con-
nate for about % their length, 7—12 x 0.75—1 mm, pale yellow to cream-colored, subulate, each with 2
confluent, dorsally-inclined apical pores; connective thickened dorsally and prolonged dorso-basally
into an acute spur 1—2 mm long. Ovary superior or 1/4- to 1/5-inferior, glabrous, 6-locular, elevated
apically into a cone about 2 mm high and a shallow collar ca. 0.5 mm high, the latter disappearing as
the ovary cone enlarges with age on mature fruits. Style glabrous, 1.2—1.4 cm long; stigma + truncate
but not conspicuously expanded. Berry becoming red at maturity on some plants, 6-9 x 7-8 mm.
Seeds | mm long, tan, narrowly ovoid.

DISTRIBUTION AND PHENOLOGY. — Locally common in rainforests from central Costa Rica
south through Panama to eastern Colombia and southeastern Venezuela, east to Surinam and French
Guiana and south to the Brazilian Amazon (Amapa, Para, Amazonas) from sea level to 1375 m.
Flowering material has been collected from March through August, October and December, fruiting
collections have been gathered from March through December.

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela: between Cataratas and
San Lorenzo about 13—17 km N of La Balsa de San Ramon, 8 June 1976, Utley & Utley 5109 (CAS,
DUKE). Puntarenas: ca. 1.1 km N of Las Cruces Tropical Botanical Garden on road to San Vito, 14
July 1977, Almeda et al. 3058 (CAS, CR). San José: Reserva Biologica Carara Estacion Bijagual,
9°46'N, 84°36'W, 23 Jul. 1990, Bello 2357 (CAS, INB, MO). PANAMA: Canal Area: Pipeline Road,
approx. 15 km N of Gamboa, 8 Oct. 1982, Schmalzel & Moreno 1100(CAS). Chiriqui: 7.5 miles from
bridge over Rio Chiriqui Viejo on road to Rio Sereno, 7 Apr. 1979, Hammel et al. 6877 (CAS, MO,
PMA). Coclé: La Mesa at El Valle de Anton, 6.4 km along La Mesa road from El Valle main road, 28
Apr. 1977, Folsom & Butcher 2834 (CAS, MO). Colon: upstream from bridge over Rio Guanche, 27
May 1980, Antonio 4793 (CAS, MO). Comarca de San Blas: Rio Playon Chico, 09°13.5’N, 78°15'W,
8 June 1994, Herrera et al. 1586 (CAS, MO). Darién: S of Garachiné on W slope of Serrania Sapo,
above place called Casa Vieja, 25 May 1991, McPherson et al. 15377 (CAS, MO, PMA). Herrera: W
of Las Minas, on Montoso de Chepo, vicinity of Chepo, 07°42'N, 80°51'W, 20 May 1987, McPherson
10931 (CAS, MO, PMA). Los Santos: Los Taretos, 10 Aug. 1962, Dwyer 2432 (MO). Panama: El
Llano-Carti highway, about 10 km N of El Llano, 20 Apr. 1973, Dressler 4360 (MO).

DISCUSSION. — The T. parasitica complex, as interpreted here, is defined by a number of consis-
tent qualitative characters. These include floral bracts that are free to the base, anther thecae that are
laterally connate for about 3/4 their length, spurlike dorso-basal anther appendages, a 6-locular ovary,
and a superior or nearly superior ovary. The morphology of hair types comprising the indument is also
diagnostic but indument presence and abundance vary considerably from population to population.
Perhaps because of its comparatively widespread geographical distribution and elevational range, T.
parasitica presents some puzzling variation in characters that are typically constant in other species.
The most conspicuous variation involves leaf size and thickness, petiole length, number of flowers per
leaf axil, peduncle length, floral bract size, calyx lobe development, hypanthial length, flower size,

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 539

and petal color. Extreme expressions for all of these characters are impressive. Previous students of
the Melastomataceae who worked with geographically defined representatives of Topobea have
seized on some of these differences in describing new taxa. Whena large number of collections are ex-
amined from across the range of this species, however, it becomes clear that much of the character
variation noted above is continuous or insignificant for the recognition of meaningful taxa. Gleason
(1958), in his treatment of Melastomataceae for Flora of Panama, recognized 7. membranacea, T.
praecox, and T. regeliana, all of which I here relegate to the synonymy of T. parasitica, the oldest
name applicable to this complex. Gleason used the name 7. praecox for plants with triangular-ovate
calyx lobes and flowers appearing when the leaves are lacking or at least not fully expanded. He de-
scribed 7. membranacea and T. regeliana as having calyx lobes that are almost obsolete and flowers
appearing while the leaves are fully expanded. Gleason further differentiated these two species by the
following couplet:

Leaves thinly coriaceous, flowers 1—2 per axil on pedicels 3-6 mm long .................... T. regeliana
Leaves membranaceous; flowers 4-6 per leaf axil on pedicels 10-15 mm long .............. T. membranacea

Unfortunately most collections attributable to this complex are in fruit, so it is difficult to criti-
cally evaluate degree of leaf maturation with flowers at anthesis. Nevertheless, the taxonomic utility
of this purported correlation seems dubious. All of the other characters that Gleason used to distin-
guish these three taxa are too variable or exhibit much overlap. Thus the character correlations used to
differentiate what he interpreted as three species make little sense when applied to the many more col-
lections now available for study from Central America. Because of this, I see no recourse other than to
recognize 7. parasitica.as a variable complex with some modally extreme character combinations that
break down as sample size increases.

In his treatment of 7. parasitica for the Flora of the Guianas, Wurdack (1993) notes that other per-
haps synonymous taxa in this complex include 7. rupicola Hoehne of Brazil and T. floribunda
Gleason and 7. rhodantha Uribe of Colombia. For illustrations of 7. parasitica see Gleason
(1958:246) and Wurdack (1993:299).

18. Topobea parvifolia (Gleason) Almeda, Proc. Calif. Acad. Sci. 46:322. 1990.

Blakea parvifolia Gleason, Phytologia 3:357. 1950. TYPE. — PANAMA. Coclé: crest of Cerro Pajita, El Valle
de Anton, 1100 m, Allen 3761 (holotype: NY!; isotype: MO!).

Tree 3—10 m tall. Uppermost branchlets glabrous, quadrisulcate but becoming rounded-quadrate
with age. Young vegetative buds and floral peduncles sparingly covered with a caducous scurfy
indument. Mature leaves of a pair essentially equal in size, adaxially glabrous, abaxially glandu-
lar-lepidote; petioles 0.6—1.4 cm long; blades coriaceous, 2.5—4.7 cm long and 1.4—2.6 cm wide, el-
liptic to obovate, the apex rounded to a blunt apiculum, the base acute to cuneate, the margin entire,
3—5-nerved with inconspicuous pustulate swellings that become perforated domatia at the abaxial
blade base in the angles between the median vein and each of the two proximal lateral veins, the trans-
verse secondary veins spaced 0.5—1 mm apart at the widest portion of the blade. Flowers erect, soli-
tary in uppermost leaf axils; peduncles 1.7—3.1 cm long. Floral bracts free and entire; outer bracts 3.5
1.5—2 mm, narrowly ovate to ovate-lanceolate, apex rounded; inner bracts 3 x 1.5—2 mm, oblong,
apex rounded. Calyx tube (fruiting hypanthia) 0.5—0.75 mm long, campanulate to cupulate; free por-
tions of calyx lobes 0.75—1 mm long and 1.5—2 mm wide basally between interlobe sinuses, broadly
triangular and often becoming bluntly apiculate on the most mature berries. Petals 6, obovate, gla-
brous with an irregular scattering of hyaline disc-shaped glands when dry, 0.7—1 x 0.5—0.6 cm, white
or white flushed with pink along the margins. Stamens 12; filaments 2.5 mm long, declinate, gla-

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brous; anthers oblong, laterally connate in a semicircular ring, 2—2.5 mm long, 0.5 mm wide, each an-
ther sac opening by a separate dorsally inclined terminal pore; connective simple. Ovary 2/3-inferior,
4-locular, glabrous at the apex which is elevated into a low truncate cone that becomes increasingly
rounded in fruit. Style glabrous, 6.5 mm long; stigma punctiform. Berry 6—6.5 x 6—6.5 mm. Seeds
1 mm long, beige, cuneate-clavate to narrowly pyriform.

DISTRIBUTION AND PHENOLOGY. — Locally common in ridgetop elfin forest on Cerro Pajita,
Cerro Gaital, and vicinity in central Panama at 900-1100 m. Flowering has been recorded in Novem-
ber and December, February, and July, fruiting collections have been made in February and July.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Cocleé: Cerro Gaital, east slope and
ridges leading to the summit, 24 Feb. 1988, Almeda et al. 5898 (CAS, PMA); vicinity of La Mesa, be-
yond El Valle, on eastern ridge along trail to summit of Cerro Gaital, 08°37'N, 80°07'W, 13 July 1987,
McPherson 11261 (CAS, MO, PMA).

DISCUSSION. — This Panamanian endemic is known only from windswept slopes and ridges in
the vicinity of El Valle de Anton. Among the species with small glabrous leaf blades and free floral
bracts, 7. parvifolia stands out by its inconspicuous pustulate swellings that become perforated
domatia at the abaxial blade base in the angles between the median vein and each of the two proximal
lateral veins. The anther thecae are unappendaged, laterally connate in a semicircular ring with each
anther sac opening by a separate terminal pore, and the ovary is 4-locular.

19. Topobea pittieri Cogn., DC. Monogr. Phan. 7:1088. 1891.

TYPE. — COSTA RICA. La Palma, alt. 1550 m, 18 Dec. 1888, Pittier 706 (holotype: BR!;
isotypes: BR!, CR!).

Terrestrial or epiphytic shrub 1-4 m tall. Uppermost branchlets obscurely rounded-quadrate be-
coming rounded with age. Young internodes, young vegetative buds, bracts, hypanthia and calyx
lobes sparsely to moderately squamulose with inconspicuous scales that may superficially appear like
sessile glands. Mature leaves of a pair equal to slightly unequal in size; petioles 1.1—2.7 cm long;
blades subcoriaceous, 6.5—14.5 x 2.6—7.5 cm, elliptic, the apex acuminate to caudate-acuminate, the
base acute to obtuse and somewhat decurrent on the petiole, the margin obscurely undulate-crenulate,
5-plinerved with an additional inconspicuous intramarginal pair, the innermost primary veins diverg-
ing from the median vein 0.3—1 cm above the blade base (on abaxial surface) with pocket domatia
formed where innermost primaries diverge from the median vein, the transverse secondary veins
spaced 0.5—1.5 mm apart at the widest portion of the blade. Flowers erect, borne in clusters of 2 to 5 in
each leaf axil of distal branches; peduncles |—2.5 cm long. Floral bracts free and much shorter than the
hypanthium and calyx; outer bracts 1.5—2.5 x 1.5—2.5 mm, ovate to deltoid; inner bracts 1—1.5 x
1.2—1.5 mm, broadly ovate to suborbicular. Hypanthium (at anthesis) narrowly campanulate, 3—3.5 x
33.5 mm. Calyx tube | mm long; calyx lobes 0.5 2-3 mm, truncate to retuse with inframarginal
teeth positioned medially on the abaxial surface. Petals 6, glabrous, 7—8.5 x 3.5—4.5 mm, white or
greenish-white, oblong-ovate and acute apically. Stamens 12; filaments 2.5—3.5 mm long, declinate
and glabrous; anthers free, 2.5—-3 x 0.5 mm, yellow, subulate, each opening by a solitary, dor-
sally-inclined pore; connective thickened dorsally and prolonged dorso-basally into a + horizontal
toothlike appendage 0.5 mm long. Ovary 3/4-inferior, 4-locular, slightly elevated at the summit into a
low glabrous laterally lobulate blunt cone surrounding the stylar scar. Style glabrous, 5—8 mm long,
stigma punctiform. Berry 5—9 x 6-8 mm, pink-purple at maturity. Seeds narrowly ovoid to cuneate,
1 mm long, deep purple with a smooth testa.

DISTRIBUTION AND PHENOLOGY. — Locally common in very wet cloud forests from Costa Rica
south to Colombia and Ecuador at 500—1800m. Flowering and fruiting specimens have been collected
during every month of the year.

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 54]

REPRESENTATIVE SPECIMENS EXAMINED. — COSTA RICA. Alajuela: about 9.7 km N of Los
Angeles de San Ramon, 23 Feb. 1978, Almeda & Nakai 3846 (CAS, CR). Cartago: Refugio Nacional
de Fauna Silvestre Tapanti, 22 Mar. 1986, Almeda et al. 5709 (CAS, CR). Heredia: 12-15 km SW of
Horquetas; Finca Rara Avis, transect SE across Rara Avis boundary along Rio Sardinal to S corner of
property on boundary with Braulio Carrillo Park, 10°20'N, 84°02'W; 20 Apr. 1988, Hammel et al.
16701 (CAS, CR, INB, MO). Limon: Canton de Talamanca, Parque Nacional Cordillera de
Talamanca, Rio Coén, entre Ujarras y San José Cabécar, 09°24'20, 83°13'30”°W, 3 Apr. 1993, Herrera
6194 (CAS, CR, INB, MO). San José: ca. 21 km N of San Isidro de Coronado on lower W slopes of
Volcan Irazu, 5 July 1977, Almeda et al. 2918 (CAS, CR). PANAMA. Bocas del Toro; Edwin Fabrega
Dam and Reserve in Fortuna along the Continental Divide trail, 12.9 km N of Sitio de Presa offices
above the dam, 08°48'04”N, 82°15'04°W, 7 Feb. 1996, Almeda et al. 7539 (CAS, PMA). Chiriqui:
Fortuna Dam area about 3.7 km S of the Sitio de Presa offices. Trail through Quebrada Aleman, 9 Feb.
1996, Almeda et al. 7553 (CAS, PMA); Cerro Colorado, Bocas Road, 17—18 Feb. 1977, Folsom et al.
1770 (CAS).

DISCUSSION. — This species is distinguished by the combination of pocket domatia on abaxial
foliar surfaces, solitary anther pores, apically acute petals, seeds that are deep purple at maturity, anda
4-locular ovary. Cogniaux (1891), Standley (1938), and Wurdack (1980) overlooked the presence of
foliar domatia in this species. He described them as “inner primaries calloused at the base beneath.”

20. Topobea pluvialis Standl., Field Mus. Nat. Hist., Bot. Ser. 22:162. 1940.

TYPE. — PANAMA. Darién: Chepigana District, crest of Cana-Cuasi trail, 15 Mar. 1960, Terry
& Terry 1560 (holotype: F!; isotype: MO!).

Reportedly a tree 4—6 m tall or an epiphytic shrub. Uppermost branchlets quadrate becoming
rounded with age and irregularly inflated into clavate-thickened formicaria just below some nodes.
Young vegetative buds and young floral peduncles caducously covered with a furfuraceous indument
of minute branlike hairs, otherwise glabrous throughout. Mature leaves of a pair somewhat unequal in
size, glabrous on both surfaces at maturity; petioles 1.3—2.5 cm long; blades coriaceous, 10—15.5 =
3.4—7.7 cm, elliptic-oblong to elliptic-obovate, the apex caudate-acuminate, the base acute to obtuse,
the margin entire, 5-plinerved with innermost primaries diverging from the median vein 3—9 mm
above the blade base (on abaxial surface) and with hair tuft domatia that are sparsely covered with
barbellate hairs in the angles formed with the median vein, the transverse secondary veins spaced
0.5—1.5 mm apart at the widest portion of the blade. Flowers erect, solitary or borne in clusters of 2—3
in each leaf axil of distal branches; peduncles 0.8—1.7 cm long. Floral bracts essentially glabrous at
maturity; outer bracts 4—7 x 4 mm, free, ovate to elliptic-lanceolate; inner bracts 3-4 x 3-4 mn, free,
depressed-ovate to suborbicular. Hypanthium cupulate, 5—6 x 6 mm. Calyx tube 2.5—3 mm long,
erect; calyx with broadly depressed rounded-triangular lobes | x 3 mm, each of which is terminated
by a tooth 0.5 mm high or the calyx often appearing nearly truncate. Petals 6, glabrous, 1—1.3 =
0.7—0.9 cm, white or pink (fide Hartman 12381), obovate, the apex bluntly acute to subrotund, the
margin entire. Stamens | 2; filaments 6-7 mm long, declinate, complanate, glabrous; anthers laterally
coherent for about 2/3 of their length, 4-5 x 0.75 mm, yellow, oblong to oblong-subulate, each with 2
confluent, dorsally-inclined apical pores; connective somewhat thickened dorsally and prolonged
dorso-basally into a deflexed tooth-like appendage 0.5—1 mm long. Ovary 1/3-inferior, 6-locular, ele-
vated apically in a short stipe (0.5 mm high) bearing the stylar scar but lacking a collar. Style glabrous,
1.3-1.4 cm long; stigma + conic when expanded. Immature berry 6 x 4 mm. Seeds mostly | mm long,
beige, the testa smooth.

542 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DISTRIBUTION AND PHENOLOGY. — A rare and little-collected species of the rainforests of
southern Panama near the Colombian frontier at 900—1650 m. Flowering collections have been made
in December and February through April, fruiting collections in April, November and December.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Darién: top of ridge leading to Cerro
Pirre, near Rancho Plastico, 13 Nov. 1977, Folsom et al. 6282 (CAS, MO); SW ridge leading to
Alturas de Nique, on border with Colombia, 29 Dec. 1980, Hartman 12381 (CAS, MO, PMA); Parque
Nacional Darién, Cerro Pirre, 7 Feb. 1991, Herrera et al. 886 (CAS, MO).

DISCUSSION. — The foliar acarodomatia of 7: pluvialis take the form of invaginated depressions
on the abaxial surface but they are inflated and paired on the adaxial surface. These domatia are lack-
ing in T. inflata Triana, a closely related Colombian species. This latter species also has fusiform
internodal swellings (presumably inhabited by ants) that are larger and more consistently developed
on uppermost internodes than in 7. pluvialis. Herrera et al. 1485 (CAS) from San Blas, Panama
(350-480 m) could represent an outlying population of 7. pluvialis or a closely related taxon but it is
in young bud and has abaxial foliar domatia that appear to be ruptured and devoid of hairs rather than
open and covered with a few barbellate hairs. Better material is needed before this entity can be identi-
fied with certainty. Topobea pluvialis is the only known member of the genus in Mesoamerica that
produces both cauline formicaria (presumably for ants) and foliar domatia for mites.

21. Topobea standleyi L. O. Williams, Fieldiana, Bot. 29:583. 1963.

TYPE. — GUATEMALA. Baja Verapaz: dry rocky hills in forest of pine and oak, north of Santa
Rosa, 30 Mar. 1939, Standley 69709 (holotype: F!; isotype: NY).

Reportedly a terrestrial shrub or tree to 8 m tall. Uppermost branchlets + terete, the very young
internodes, vegetative buds, petioles on young leaves, floral peduncles, and abaxial leaf surfaces
moderately furfuraceous with caducous branlike or stellulate hairs. Mature leaves of a pair somewhat
unequal in size, glabrous on the adaxial surface at maturity; petioles 0.5-4 cm long; blades +
coriaceous, 9.5—15 x 5—10 cm, elliptic to elliptic-ovate, the apex abruptly acuminate, the base acute,
the margin entire, 5-nerved (the outermost pair often obscure), the transverse secondary veins spaced
0.5—1 mm apart at the widest portion of the blade. Flowers erect, solitary or paired in each leaf axil of
upper branches; peduncles 0.7—1.2 cm long. Floral bracts moderately to sparsely covered with a
lanuginose indument of caducous matted hairs at maturity; outer bracts 6-8 x 5 mm, free, ovate, typi-
cally with a keeled median vein; inner bracts 5—6 x 5 mm, free, suborbicular to obovate. Hypanthium
cupulate, 0.7—0.8 < 0.7 cm (at anthesis). Calyx tube 2.5 mm long, erect; calyx lobes bluntly deltoid
and + rounded apically (often appearing like erect blunt teeth on fruiting hypanthia), 1.5—2 x 2 mm,
covered with a brown lanuginose indument of woolly matted hairs. Petals 6, glabrous, 0.7—1.1 x
0.5—0.9 cm, pink and white (fide Standley 69709), obovate, the margin obscurely retrorse-ciliolate.
Stamens 12; filaments 5 mm long, declinate, complanate, glabrous; anthers free, 5.5—7 x 1 mm, ob-
long, each with 2 divergent, dorsally-inclined apical pores; connective somewhat thickened dorsally
but not elaborated into an appendage. Ovary 1/2-inferior, 6-locular, the apex smooth, glabrous and
lacking a cone and collar. Style glabrous, 9 mm long; stigma punctiform. Berry 8 x 5 mm. Seeds
1.5 mm long, beige with a black lateral raphe and smooth testa.

DISTRIBUTION AND PHENOLOGY. — Rocky hills in pine-oak forests of Central Guatemala at
1500 m. The two known collections, which are in flower and fruit, were collected in March and April.

REPRESENTATIVE SPECIMENS EXAMINED. — GUATEMALA. Baja Verapaz: pine-oak forest on
rocky hills near and above Santa Rosa, 4 Apr. 1941, Standley 91045 (F, NY).

DISCUSSION. — This species, which has not been collected since April of 1941, appears to be a
local endemic known only from the type and the one other collection cited above. It is readily recog-
nized by its abruptly acuminate leaves, caducous lanuginose indument on hypanthia, calyx lobes, and

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 543

floral bracts, and unappendaged anthers, each of which has two divergent apical pores. For an illustra-
tion of this species see Standley and Williams (1963:520).

22. Topobea suaveolens Almeda, Proc. Calif. Acad. Sci. 46:323. 1990.

TYPE. — PANAMA. Veraguas: along trail to summit of Cerro Tute about 4 mile above the
Escuela de Agricultura Alto Piedra near Santa Fe, 29 Jan. 1989, Almeda et al. 6484 (holotype: CAS!;
isotypes: AAU!, BM!, BR!, CR!, DUKE!, F!, MEXU!, MICH!, MO!, NY!, PMA!, TEX!, US!).

Epiphytic tree to 4 m tall, often obscuring and overtaking the crowns of host trees. Uppermost
branchlets quadrate to quadrisulcate and glabrous with well-defined interpetiolar ridges or lines. Veg-
etative buds caducously lepidote-furfuraceous. Mature leaves of a pair equal or slightly unequal in
size, glabrous throughout; blades subcoriaceous, 3.6—5.3 x 1.6—2.5 cm, elliptic, the apex acuminate to
caudate-acuminate, the base acute, the margin entire, 3-plinerved with an additional inconspicuous
pair of submarginal veins, transverse secondary veins spaced mostly 0.25 mm apart at the widest por-
tion of the blade, pocket domatia typically formed abaxially in the angles between the median vein and
the two proximal lateral veins. Flowers pendent and solitary in the leaf axils of uppermost branches;
peduncles 2.3—3 cm long, glabrous. Floral bracts glabrous, entire; outer bracts 4.5—7.5 x 3-4 mm, el-
liptic to elliptic-ovate, apex bluntly acute to rounded; inner bracts 44.5 x 4mm, ovate to
suborbicular, apex broadly rounded. Calyx tube 1.5 mm long; calyx lobes | mm long and 4 mm wide
basally, broadly ovate to deltoid-ovate with a blunt callose-thickened tooth on the abaxial apex of
each lobe, margin entire, glabrous throughout. Petals 6, glabrous, connivent to somewhat imbricate
and bell-like when fully expanded, 1.2—1.5 x 0.9-1.1 cm, white flushed with dark pink along a broad
marginal band, obovate, apically rounded, entire. Stamens 12, free and encircling the exserted style;
filaments glabrous, 2.5—-3.5 mm long; anthers 2.5 mm long, | mm wide, yellow, laterally com-
pressed, oblong in ventral view and narrowly ovoid in profile view with a shallow dorso-basal depres-
sion at the filament insertion, tipped with a solitary, dorsally-inclined pore 0.75 mm in diameter;
connective unappendaged. Ovary 1/2-inferior, 6-locular, elevated at the glabrous apex into a smooth
dome |—1.5 mm high. Style straight, glabrous, 8.5—9 mm long; stigma truncate. Berry 5—6 x 6—7 mm.
Seeds narrowly and irregularly pyriform, 0.75—1 mm long.

DISTRIBUTION AND PHENOLOGY. — Known only from the cloud forests of Cerro Tute in
west-central Panama at 850—1100 m. The three known collections, all of which are in flower and fruit,
were collected from January through March.

REPRESENTATIVE SPECIMENS EXAMINED. — PANAMA. Veraguas: vicinity of Cerro Tute,
along trail to summit, 08°30’N, 81°07'W, 19 Mar. 1987, McPherson 10654 (CAS, MO, PMA); near
Cerro Tute-Arizona, above Santa Fe and Alto de Piedra, 8°30'N, 81°10'W, 5 Feb. 1988, McPherson
12043 (CAS, MO, PMA).

DISCUSSION. — This little-collected Panamanian endemic appears to be restricted to the slopes
of Cerro Tute. It is unusual among congeners in having sweet-smelling flowers that are pendent with
free connivent anthers that form a ring around the exserted style. Topobea suaveolens also has distinc-
tive laterally compressed anthers that are uniporose and narrowly ovate in profile view with a conspic-
uous dorso-basal depression near the filament insertion. For an illustration of this species see Almeda
(1990:324).

23. Topobea tetramera Almeda, sp. nov. (Fig. 5)

TYPE. — PANAMA. Veraguas: headwaters of Rio Caloveborita ca. 15 km past Escuela
Agricola Alto Piedra above Santa Fé on the Atlantic watershed, 500 m, 16 May 1981, Sytsma & An-
derson 4758 (holotype: CAS!; isotype: MO, PMA).

544 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Frutex epiphyticus vel terrestris. Ramuli sicut pedunculi folia inflorescentia hypanthiaque pilis
1—2 mm longis induti. Folia in quoque pari dimorpha papyracea et denticulata. Folia maiora: lamina
4.5-9.4 x 2.3-4.3.cm elliptica vel elliptico-ovata vel elliptico-obovata apice acuminata basi
asymmetrice obtusa vel rotundata 5-plinervata. Folia minora: lamina 0.7—1.7 x 0.6—1 cm ovata apice
acuta basi obtusa vel rotundata 3-nervata. Flores 4-meri in quoque nodo superiori singuli; pedunculis
3-4 mm longis; bracteae omnino liberae 0.7 x 0.2—0.4 cm elliptica apice acuta; bracteae interiores
omnino liberae 0.9 x 0.3—0.4 cm elliptica apice acuta vel acuminata. Calycis tubus | mm longus, lobis
0.7 x 0.2 cm. Petala 0.8—0.9 = 0.5—0.7 cm obovata. Antherarum thecae 2 x 0.75 mm inter se non
cohaerentes, dorsaliter biporosae; connectivum nec prolongatum nec appendiculatum. Ovarium
4-loculare et omnino inferum, cono glabro (collo non evoluto).

Epiphytic or terrestrial shrub to 2 m tall. Uppermost branchlets quadrate becoming rounded with
age. Cauline internodes, leaf blades (both surfaces), peduncles, floral bracts, hypanthia, and calyx
lobes moderately to copiously hirsute with rusty brown hairs mostly 1-2 mm long. Mature leaves ofa
pair markedly unequal in size; blades coarsely papery when dry, the larger one at each node 4.5—9.4 x
2.34.3 cm, elliptic varying to elliptic-ovate or elliptic-obovate, the apex abruptly acuminate, the base
somewhat obliquely obtuse to rounded, the margin denticulate (sometimes obscurely so), 5-plinerved
with the innermost pair of primary veins diverging from the median vein 2—5 mm above the blade
base, the transverse secondary veins spaced 24 mm apart at the widest portion of the blade; petiole
5—7 mm long; the smaller blade 0.7—1.7 x 0.6—1 cm, ovate, apex acute, base rounded to obtuse, mar-
gin entire, 3-nerved, the transverse secondary veins not evident; petiole 2—3 mm long. Flowers erect,
solitary in each axil of uppermost leaves, peduncles 3—4 mm long. Floral bracts green and entire; outer
bracts 0.7 x 0.2—0.4 cm, free, narrowly elliptic, apex acute; inner bracts 0.9—1 x 0.3—0.4 cm, free, nar-
rowly elliptic, apex acute to acuminate. Hypanthium (at anthesis) 4 mm long to the torus and 4 mm in
diameter. Calyx tube | mm long, erect or somewhat flaring at anthesis. Calyx lobes (at anthesis) 4,
deltoid at the base but abruptly tapered to narrow linear segments 0.7 cm long and 0.2 cm wide at the
base between sinuses. Petals 4, glabrous, 0.8—0.9 x 0.5—0.7 cm, translucent white (fide de Nevers et al.
5460) or translucent pink (fide Sytsma & Andersson 4758), thin and translucent when dry, obovate,
the apex rounded, the base broadly clawed, entire. Stamens 8, isomorphic; filaments 3—3.5 x 0.5 mm,
complanate, glabrous; anthers free, 2 x 0.75 mm, yellow, narrowly ovoid, each with 2 confluent,
somewhat dorsally-inclined pores at the truncate apex; connective thickened dorsally but
unappendaged. Ovary completely inferior (at anthesis), 4-locular, glabrous at the summit which is el-
evated into a blunt cone surrounding the stylar scar. Style glabrous, 5.5-6 mm long; stigma
punctiform. Berry red at maturity, 6-8 x 7-8 mm. Seeds | mm long, white, narrowly pyriform to
cuneate, testa smooth.

DISTRIBUTION AND PHENOLOGY. — Local and evidently uncommon in low rainforests from the
Fortuna region of western Panama east to Cerro Brewster in central Panama at 800—1100 m. Collected
in flower in April and May, in fruit in November.

ADDITIONAL SPECIMENS EXAMINED. — PANAMA. Chiriqui: Fortuna Dam area, along
Quebrada Bonito to E of road, 8°45'N, 82°13’W, 8 Feb. 1984 (sterile), Churchill et al. 4767 (MO).
Comarca de San Blas: Cerro Brewster, premontane rain forest, 9°18'N, 79°16’W, 21 Apr. 1985, de
Nevers et al. 5460 (CAS); Cerro Brewster, premontane rain forest, 9°18'N, 79°16'W, 20 Nov. 1985, de
Nevers et al. 6266 (CAS). Veraguas: 11 km from Escuela Agricola Alto de Piedra, along Rio Dos
Bocas, 15 Nov. 1974, Mori & Kallunki 3092 (CAS, MO).

DISCUSSION. — This extraordinary species appears to be highly derived because of its 4-merous
flowers (Fig. SF) with eight stamens and a 4-locular ovary. The strongly dimorphic leaves at each
node (Fig 5A) and the copious rusty brown hirsute pubescence on vegetative and floral parts are also
characteristic. These latter features are shared with 7. dimorphophylla which differs in having partly
fused inner floral bracts, 6-merous flowers with 12 stamens, and a 6-locular ovary.

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 545

FIGURE 5. Topobea tetramera Almeda. A. habit, 1/5; B. foliar dimorphism at a node, ca. 1; C. fruiting hypanthium with at-
tached decussate floral bracts, ca. 3; D. outer floral bract (adaxial surface), ca. 3; E. inner floral bract (adaxial surface), ca. 3;
F. hypanthium (at anthesis) with floral bracts, petals, androecium, and style removed, ca. 3; G. petal (adaxiall surface), ca. 3;

H. stamens, % profile view (left) and dorsal view (right), 10;1. seeds, 20.(A,B from Mori & Kallunki 3092; C—1 from Sytsma
& Anderson 4758.)

546 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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ETYMOLOGY. — The epithet for this species is derived from the Greek words tetra, meaning
four, and merus, referring to number of parts, in reference to the unusual 4-merous flowers and
4-locular ovary of this species.

24. Topobea watsonii Cogn., DC. Monogr. Phan. 7:1089. 1891.
TYPE. — GUATEMALA. Prope Chocon River, 11 Mar. 1885, Watson 94/211 (holotype: BR!).

Topobea rosea Gleason, Publ. Carnegie Inst. Wash. 522: 536. 1940. TYPE. — BELIZE. Temash River, 13 Mar.
1936, Schipp 1320 (holotype: NY!)

Topobea urophylla Standl., Field Mus. Nat. Hist., Bot. Ser. 22:162. 1940. TYPE. — PANAMA. Darién:
Chepigana District, Rio Balsa, above Tucuti, 6 Mar. 1940. Terry & Terry 1411 (holotype: F!; isotype: MO!).

Topobea cooperi Gleason, Phytologia 3:354. 1950. TYPE. — PANAMA. Bocas del Toro: Cricamola Valley,
Region of Almirante, Jan.—Mar. 1928, Cooper 199 (holotype: NY!; isotype: F!).

Topobea allenii Standl. & L. O. Williams, Ceiba 3:216. 1953. TYPE. — COSTA RICA. Puntarenas: Esquinas
Forest, region between Rio Esquinas and Palmar Sur de Osa, alt. 75 m, 5 Feb. 1951, Allen 5844 (holotype:
EAP; isotypes: F!, US!).

Scandent epiphytic shrub with lax branches 3—4 m long. Uppermost branchlets bluntly quadrate
but becoming rounded with age. Juvenile growth, uppermost cauline internodes, peduncles, and floral
bracts moderately to densely covered with barbellate or stellulate and scurfy hairs but glabrate with
age. Mature leaves of a pair equal to somewhat unequal in size, adaxially glabrous, abaxially sparsely
covered with plumose or barbellate hairs (mostly on the elevated primary veins) and stellulate or
branlike hairs (especially on the actual surface) varying to nearly glabrous; petioles |—4.8 cm long;
blades coriaceous, 5.5—16.5 x 3—7.7 cm, elliptic to elliptic-ovate, the apex long-acuminate to abruptly
caudate, the base obtuse to rounded, margin entire, 3—5-nerved or 3—5-plinerved (sometimes with an
intramarginal pair of obscure veins), the transverse secondary veins spaced 0.25—0.5 mm apart at the
widest portion of the blade. Flowers erect, |—4 per leaf axil of uppermost branches; peduncles
0.7—1.3 cm long. Floral bracts free or connate basally for 1-2 mm, elliptic to elliptic-ovate, apex acute
to rounded apically; outer and inner bracts 0.6—1 = 0.4-0.8cm. Calyx tube 3-4 mm long,
campanulate; free portions of calyx lobes 2-3 mm long and 2.5—3.5 mm wide basally between
interlobe sinuses, bluntly triangular with a prominently elevated callose-thickened tooth or append-
age at the median apex abaxially. Petals 6, glabrous or stellulate puberulent abaxially, 10-16 x
4.5—7 mm, pink, rhombic-ovate to obovate, the apex + acute. Stamens 12; filaments 5—7 mm long,
declinate, complanate and glabrous; anthers laterally connate, 6—7.5 x 1 mm, yellow, granulose along
lower ventral half of the thecae, each with 2 confluent, dorsally-inclined pores; connective prolonged
dorso-basally into a deflexed caudiform appendage 0.5—1 mm long. Ovary 1/3-inferior, 6-locular,
glabrous and elevated at the apex into a lobulate stylar collar |—1.5 mm high. Style glabrous,
1.2—1.6 cm long; stigma capitellate. Berry red, |—1.3 x 0.9-1.3 cm. Seeds |—1.5 mm long, beige, nar-
rowly ovoid to cuneate.

DISTRIBUTION AND PHENOLOGY. — Locally common in low rainforests and cloud forests from
southern Mexico (Chiapas) and Belize south through Central America (excluding El Salvador) to Co-
lombia from sea level to 1400 m. Flowering specimens have been collected from October through
February and in May, fruiting collections from February through September.

REPRESENTATIVE SPECIMENS EXAMINED. — BELIZE. Toledo: Edwards Road beyond Colum-
bia, on high ridge, 1 1 Nov. 1947, Gentle 6316 (CAS, LL). COSTA RICA. Puntarenas: between Golfo
Dulce and Rio Térraba, Nov. 1947 (without exact date), Skutch 5263 (US). GUATEMALA. Petén:
Los Arcos, 2 km, E of Km 143 on Cadenas Road, 17 Dec. 1969, Contreras 9379 (CAS, LL, US).
Izabal: Rio Chacon, 8 Feb. 1921, Johnson 1237 (US). HONDURAS. Gracias a Dios: Ahuas Bila,
200 km SO de Puerto Lempira, orilla del Rio Wanki, Coco 0 Segovia, 5—13 May 1985, Nelson & Cruz

ALMEDA: TOPOBEA IN MEXICO AND CENTRAL AMERICA 547

9319 (CAS, TEFH). MEXICO. Chiapas: Municipio Las Margaritas, low ridges at the confluence of
the Rio Lacantum (Rio Jataté) on the Guatemala border, 14 Mar. 1973, Breedlove & McClintock
34103 (DS, MEXU). NICARAGUA. Jinotega: below Penas Blancas via El Tuma, 6 May 1976, Neil
253 (CAS, MO). Matagalpa: Macizos de Pefias Blancas, SE side, drainage of Quebrada El
Quebradon, ca. 13°14—15'N, 85°38'W, 20-21 Jan. 1982, Stevens et al. 21282 (CAS, MO). Nueva
Segovia: 10 km SE of Jalapa, 24 Dec. 1973, Atwood et al. 6827a (CAS, MO). Zelaya: along road from
Bonanza (airstrip) through Constancia (mineshaft) to Laguna Siempreviva (dam), 23 Apr. 1978,
Stevens 8015 (CAS, MO). PANAMA. Bocas del Toro: Laguna de Chiriqui, 15 km oeste de Punta
Cricamola, entrando Ensenada de Catavela, y subiendo Quebrada Nuri, 8°55'N, 81°49'W, 19 Mar.
1993, Foster et al. 14602 (CAS, PMA). Coclé: Caribbean side of divide at El Copé, 80°35’ W, 8°45'N,
4 Feb. 1983, Hamilton & Davidse 2769 (CAS, MO, PMA). Colon: lumber road on Santa Rita east
ridge, 23 Feb. 1968, Correa & Dressler 748 (MO). Comarca de San Blas: trail from Puerto Obaldia in-
land towards Bongo, 24 Mar. 1985, D’Arcy & McPherson 16142 (MO): Playon Chico, Rio Ukupseni
caminando por el Rio Ukupseni, 09°15’N, 78°15'W, 30 Oct. 1991, Herrera et al. 1024 (CAS, MO,
PMA). Darién: Parque Nacional del Darién, ridge between N & S branches of Rio Pucuro, across river
from old Kuna village of Tacarcuna ca. 18 km E of Pucuro, 8°04'N, 77°16'W, 21 Oct. 1987, Hammel
etal. 16336(CAS, MO, PMA). Panama; along newly cut road from El Llano to Carti-Tupile, 12 miles
above Pan-American highway, 13 Mar. 1973, Croat 22899 (CAS).

DISCUSSION. — This species is defined by a number of diagnostic characters. The petals are
thombic-ovate to obovate and stellulate-puberulent abaxially (in part), the anthers are dorso-basally
appendiculate, the thecae are laterally coherent for about 2/3 of their length and distinctly granulose
along their lower ventral sides, and the ovary has a conspicuous fimbriate-lobulate stylar collar. This
comparatively widespread species is variable in the density and persistence of the indument on
cauline internodes, peduncles, and floral bracts. It is otherwise uniform in all other diagnostic fea-
tures. The types of 7. allenii, T. cooperi, T. rosea, and T. urophylla, taxa here relegated to synonymy,
are good matches for typical 7. watsonii in all diagnostic features. When Standley and Williams de-
scribed T. allenii from Costa Rica they suggested a relationship with 7. urophylla which is under-
standable in view of the taxonomy adopted here. Gleason also appears to have been unaware of T.
watsonii when he described T. rosea from Belize because he compared it only to 7. calycularis in the
protologue. In his treatment of Topobea for Flora of Panama, Gleason (1958) recognized only 7.
cooperi and T. urophylla. His key to the genus suggests that he maintained these species based on a
sparse furfuraceous indument for the former and an indument of barbellate hairs for the latter. Speci-
mens from throughout the range of this species often have varying combinations and densities of these
two indument types making Gleason’s purported distinctions taxonomically insignificant.

ACKNOWLEDGMENTS

Field work for this study was made possible, in part, by U. S. National Science Foundation Grants
DEB 76-83040, DEB 78-25620, BSR 86-14880 (Flora Mesoamericana), the G. Lindsay Field Re-
search Fund and the Nathan Jay and Virginia Friedman Fund of the California Academy of Sciences,
and Dr. and Mrs. Benjamin Hammett. A number of organizations provided logistical support during
field studies in Costa Rica and Panama over many years. I am especially grateful to the Museo
Nacional de Costa Rica, the Missouri Botanical Garden, the Organization for Tropical Studies, the
Instituto Nacional de Biodiversidad, Costa Rica, the Smithsonian Tropical Research Institute, and the
University of Panama. I also thank the following colleagues and friends who contributed to this study
directly or indirectly in diverse ways: J. Albright-Souza, B. Anderson, D. E. Breedlove, W. Burger,
M. Correa, T. Croat, T. Daniel, G. Davidse, G. de Nevers, C. Galdames, M. Grayum, B. Hammel, Q.
Jimenez, K. Nakai, J. Ochoa, G. McPherson, J. Solomon, D. Stevens, G. Umafia Dodero, I.
Valdespino, R. L. Wilbur, N. Zamora, and the late J. J. Wurdack. The line drawings were prepared by

548 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 52, No. 22

J. Speckels (Figures 2, 4 and 5); H. Pazdirkova (Figure 1); and S. Myers (Figure 3). The color plate
was prepared by Meg Stalcup while she was an Illustration Intern at CAS during the summer of 2001.
This internship was made possible by the Fellows of the California Academy of Sciences. I am appre-
ciative of the many courtesies received from curators and staffs of the following herbaria for either
loans, gifts of specimens, or assistance during study visits: BM, BR, C, CAS, CR, DAV, DS, DUKE,
F, G, INB, K, LL, M, MA, MEXU, MICH, MO, NY, P, PMA, SCZ, TEFH, TEX, UC, US, USJ, WIS.

LITERATURE CITED

ALMEDA, F. 1984. New and noteworthy additions to the Melastomataceae of Panama. Proc. Calif. Acad. Sci.
43(17):269-282.

. 1990. New species and new combinations in Blakea and Topobea (Melastomataceae), with an historical

perspective on generic limits in the tribe Blakeeae. Proc. Calif. Acad. Sci. 46(14):299-326.

. 2000a. The hexandrous species of Topobea (Melastomataceae). Proc. Calif. Acad. Sci. 52(9):97—109.

. 2000b. A synopsis of the genus Blakea (Melastomataceae) in Mexico and Central America. Novon

10:299-319.

. 2001. Three new Mesoamerican species of 7opobea (Melastomataceae). Brittonia 53(1):157—166.

COGNIAUX, C. A. 1891. Mélastomacées. /n Monographiae phanerogamarum, A. and C. de Candolle, eds.
7:1—1256. G. Masson, Paris.

GLEASON, H. A. 1939. The genus Clidemia in Mexico and Central America. Brittonia 3:97—140.

. 1958. Melastomataceae. /n Flora of Panama, R. E. Woodson, Jr. and R. W. Schery, eds. Ann. Missouri
Bot. Gard. 45:203-304.

LUMER, C. 2000. The reproductive biology of Blakea and Topobea (Melastomataceae). /n Monteverde: Ecology
and conservation of a tropical cloud forest, N. M. Nadkarni and N. T Wheelwright, eds. Oxford University
Press, Oxford.

STANDLEY, P.C. 1924. Melastomaceae. /n Trees and shrubs of Mexico. Contr. U.S. Natl. Herb.
23(4):1046—-1074.

. 1938. Melastomaceae. /n Flora of Costa Rica. Field. Mus. Nat. Hist., Bot. Ser. 18(3):783-845.

STANDLEY, P.C. AND L.O. WILLIAMS. 1963. Melastomaceae. /n Flora of Guatemala. Fieldiana, Bot.
24:407-S25.

WALTER, D. E. AND H. C. PRocTor. 1999. Mites: Ecology, evolution and behaviour. University of New South
Wales Press Ltd., Sydney.

WINKLER, S. 1965. Die Melastomataceae von El Salvador C. A. Bot. Jahrb. Syst. 83:33 1-369.

WURDACK, J. J. 1980. Melastomataceae. /n Flora of Ecuador, G. Harling and B. Sparre, eds. 13:1—406.

. 1993. Melastomataceae (Topobea). In Flora of the Guianas, A. R. A. Gorts-van Rijn, ed. 13:9-21.

Koeltz Scientific Books, K6nigstein.

© CALIFORNIA ACADEMY OF SCIENCES, 2001
Golden Gate Park
San Francisco, California 94118

Me

INDEX TO TAXA IN VOLUME 52PR | 1 2002

(Compiled by Hillary Culhane)

Aldisa albatrossae

Aldisa williamsi

Ceraeochrysa dislepis

Ceraeochrysa dolichosvela
Ceraeochrysa squama

Chromodoris buchananae
Chrysoperla raimundoi

Chrysopodes (Chrysopodes) adynatos
Chrysopodes (Chrysopodes) copia
Chrysopodes (Chrysopodes) crocinus
Chrysopodes (Chrysopodes) delicata
Chrysopodes (Chrysopodes) elongata
Chrysopodes (Chrysopodes) nigropicta
Chrysopodes (Neosuarius) karinae
Dimorphophyton

Eleutherobia vinadigitaria
Eleutherobia zanahoria

Eusphairica

Eusphairica distubula

Forcepia (Forcepia) acanthostylosa
Forcepia (Forcepia) elvini

Forcepia (Forcepia) hartmani
Forcepia (Forcepia) macrostylosa
Hemibagrus imbrifer

Hemibagrus variegatus
Lampophyton

Lanthanocephalus

Lanthanocephalus clandestinus
Leucochrysa (Leucochrysa) bruneola
Leucochrysa (Leucochrysa) catarinae
Leucochrysa (Nodita) affinis
Leucochrysa (Nodita) barrei
Leucochrysa (Nodita) confusa
Leucochrysa (Nodita) cornuta
Leucochrysa (Nodita) forciformis
Leucochrysa (Nodita) furcata
Leucochrysa (Nodita) guataparensis

New Taxa

Leucochrysa (Nodita) ictericus
Leucochrysa (Nodita) incognita
Leucochrysa (Nodita) interata
Leucochrysa (Nodita) lineata
Leucochrysa (Nodita) maculata
Leucochrysa (Nodita) michelini
Leucochrysa (Nodita) parallela
Leucochrysa (Nodita) retusa
Leucochrysa (Nodita) robusta
Leucochrysa (Nodita) santini
Leucochrysa (Nodita) scomparini
Leucochrysa (Nodita) squamisetosa
Leucochrysa (Nodita) tabacinus
Leucochrysa (Nodita) tenuis
Leucochrysa (Nodita) vignisi
Leucochrysa (Nodita) vittata
Lycodon zawi

Miconia colliculosa

Miconia correae

Miconia crocata

Miconia jefensis

Miconia morii

Miconia talamancensis
Miconia vestita

Nacarina aculeata

Nacarina gladius

Nacarina lavrasana

Nacarina sagitta

Paleochiton siskiyouensis
Phrynobatrachus irangi
Plesiochrysa alytos

Thairoplax merriami

Topobea amplifolia

Topobea arboricola

Topobea dimorphophylla
Topobea gerardoana

Topobea tetramera

General Index to Taxa

A

Abachrysa 251

Acanthaceae 143, 144, 152
Barlerieae 146

Haselhoffieae 143
Justicieae 146, 150
Justicieae, Diclipterinae 146, 148, 152

[549]

550 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 52

Justicieae, Justiciinae 148, 152
Justicieae, Rhytiglossinae 148
Louteridieae 143
Rhombochlamydeae 143
Ruellieae, Barleriinae 146
Ruellieae, Ruelliinae 146
Stenandriopsideae 143
Whitfieldieae 143
Acanthogorgiidae 71
Acanthopsis 152
Acanthus pubescens 155
Acari 248
Acrophytum claviger 71
Actinella 11, 18, 20
Actinocyclidae 84
Actinocyclus 84
Actinoptilum molle 224
Adelobotrys jefensis 44
Adhatoda 148
Alastega lira 28
Alcyonacea
Isididae 209
Alcyonaria 216
Alcyonacea 216
Gorgonacea 216
Pennatulacea 216
Alcyoniidae 66, 71, 88, 160, 198, 210
Alcyonium 65, 70, 72, 216, 219
digitatum 70
fauri 70
mutabiliforme 68
planiceps 66
utinomii 70
spp. 71
Aldisa 171, 175-178
albatrossae 171, 172, 176, 178, 180
barlettai 171, 176, 178
pikokai 171, 175, 178
williamsi 174, 176-178, 180
Amphibelemnon 87
Amphicampa 11, 18, 20
eruca 12, 15
Amphipleura 20
Amphora 18
Anacardium occidentale 249
Anisotes 152
Anthomastus 71
Anthothelidae 71
Aphidoidea 248
Ardeadoris 84
Arthroleptis 55
Asbestopluma 227, 242

lycopodium 227
Asteracantha longifolia 155
Asthenomacrurus 435

fragilis 435

victoris 435
Asystasia 144

coromandeliana 144

gangetica 144, 155
Aulojusticia 144, 145, 148, 152

linifolia 144, 145

B
Bagrus
corsula 130, 132
menoda 136
punctatus 137
trachacanthus 130, 132
Barleria 145, 146
cristata 145
cristata var. dichotoma 145
repens 155
senensis 145
Barleriinae 146
Bathygadidae 407, 420
Bathygadus 420, 421, 423
cottoides 417, 421-423
furvescens 417, 421, 422
spongiceps 423
sp. cf. spongiceps 417, 423
Bellonella 160, 198
spp. 71
Belonopteryx 251
Blakea 97,512
aeruginosa 516
crassifolia 105
intercepta 533, 534
micrantha \02
multiflora 535
parvifolia 539
wilburiana 530
Blakeeae 511, 512
Blepharis
boerhaaviaefolia 155
integrifolia 155
maderapatensis 155
molluginifolia 155
repens 155

C

Cadlina 78, 82, 84, 85
flavomaculata 84
limbaughorum 84

INDEX

luarna 78, 84, 85 tucumana 249, 265

luteomarginata 84 valida 260

modesta 84 spp. 248

sparsa 84 Ceratocaulon wandeli 94
Cadlinella 84 Cetonurus 458
Caelorinchus 407, 435, 445, 450, 453 crassiceps 458

acanthiger 417, 440, 447, 449, 453, 456 globiceps 417, 458

amydrozosterus 454 Chaetacanthus 152

australis 408, 414, 440, 442, 450 Chelodes 26, 28

biclinozonalis 442 raaschi 24

cingulatus 417, 442 Chileranthemum 150

fasciatus 408, 417, 443, 450, 454 Chorisochora 152

innotabilis 408, 417, 443, 445 Chromodorididae 78, 82, 84, 111, 112

kaiyomaru 417, 445 Chromodoris 112, 119

kermadecus 417, 440, 447, 453 africana 116

macrorhynchus 417, 440, 445, 447, 449, buchananae 112, 116

455, 456 decora 116

Macrourus 408 inornata 119

matamuus 417, 435, 445, 449 lekker 116

maurofasciatus 417, 443, 450, 454 magnifica 116

mirus 408, 414, 450, 452, 454 quadricolor 116

mycterismus 417, 440, 452, 453 sp. 171, 180

parvifasciatus 417, 443, 453, 454 Chrysogorgiidae 71

pseudoparallelus 440 Chrysopa 248, 250, 256, 277, 289

smithi 417, 440, 445, 449, 455 conformis 289

supernasutus 417, 440, 449, 456 oculata 248

trachycarus 417, 440, 456 paraguaria 263

sp.cf. cingulatus 442 paraquaria 263
Cavernularia 72 Chrysoperla 246, 248, 256, 267
Ceraeochrysa 256, 257, 263-265 carnea 248

acmon 258 defreitasi 249, 268, 269

berlandi 265 externa 249, 265, 268, 269, 302

caligata 249, 259 raimondoi 269

cincta 249, 257, 259 raimundoi 268

citrinus 249 Chrysoperla (Neosuarius) 270

claveri 249, 259, 260, 262 Chrysopidae 250

costaricensis 261, 265, 267 Chrysopini 250, 256, 277, 302

cubana 249, 260, 262, 263 Belonopterygini 251

dislepis 265, 267 Chrysopodes 257, 271, 272

dolichosvela 249, 266 adynatos 275

everes 249, 261, 265 adynatus 249

lineaticornis 265 collaris 276

montouana 249 crocinus 249

montoyana 259, 260, 262 delicata 249

paraguaria 249, 260, 262 divisa 249

pennyi 249 elongata 249, 275

raimundoi 249 indentata 249, 275

sanchezi 249, 263 karinae 249

scapularis 249, 259, 260, 262, 263 lineafrons 249, 275

squama 249, 261, 265, 267 nigropicta 249

tauberae 307 polygonica 249

tenuicornis 249, 264 spinella 249, 275

daz2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 52

Chrysopodes (Chrysopodes) 270, 273

adynatos 272, 273
copia 273
crocinus 273,274
delicata 274
elongata 275
indentata 273
lineafrons 270, 272
nigropicta 275
polygonica 271
spinella 272
victoriae 273
Chrysopodes (Neosuarius) 274, 276
divisa 276
karinae 276
Citrus sinensis 249
Cladorhizidae 242
Clidemia costaricensis 525
Coccoidea 248
Coelosphaeridae 228
Coleoptera 248
Coniopterygidae 248
Coryphaenoides 407, 459
carapinus 460, 464
dossenus 417, 460, 461
fernandezianus 462
filicauda 408, 417, 463, 464
grahami 417, 464, 465, 470
macrocephalus 467
memillani 465, 471
murrayi 466
paradoxus 467
quadripennatus 471
rudis 417, 467
serrulatus 417, 440, 459, 467
serrulatus oceanus 469
serrulatus serrulatus 469
striaturus 408, 417, 465, 469, 470
subserrulatus 417, 465, 470, 471
sp. cf. fernandezianus 462
Coryphaenoides (Chalinura) 470
Crabbea 146
angustifolia 146
reticulata 155
velutina 146, 155
Cucumis melo 249
Cynomacrurus 47]
piriei 414, 417, 471, 492

D

Dermatobranchus 68
Desmogonium 1]

Diaulula
sandiegensis 183-185, 190, 191
sp. #1 183
Dicliptera 146-148, 152
bupleuroides 146
cuneata 146
elegans 146
heterostegia 146
leonotis 146
magaliesbergensis 146
parvibracteata 146
roxburghiana 146
verticillata 146
Dimorphognathus 55
Dimorphophyton 68, 70, 87
mutabiliforme 68, 71
Diprionidae 248
Discodoris liturata 180
Doridigitata 183
maculata 183
Doris 184, 190
(s.1.) sp. 183-185, 187, 190
echinata 183, 190
odonoghuei 183, 184, 190, 191
sp. 183
sp. #1 171
sp. #2 180
sp. #10 174
Duosperma 152
Durvilledoris pusilla 123
Duvernoia 144, 148, 152
aconitiflora 148
adhatodoides 148
Dyschoriste depressa 155

E

Eleutherobia 70-72, 159, 160, 163, 195, 198,

209! 210: 2122152219
aurea 195, 207

grayi 163, 164, 209, 210, 212, 215, 216

japonica 160, 198, 210
lutea 159, 209

rotifera 195

rubra 207

studeri 195

vinadigitaria 198, 201, 202, 207, 209
zanahoria 160, 163, 164, 195, 209, 215

sp. 210

spp. indet. 219
Ellisellidae 71
Elytraria acaulis 155
Eopteria 28

INDEX

struszi 29
Eopteriacea 28

Eopteriidae 28
Eranthemum 146
Esperiopsis forcipula 241
Euchasuna 28
Eunophora 18, 20
Eunotia 11, 18, 20

bilunaris 20

conversa 12, 15, 18, 20

curvula 12, 18, 20

synedraeformis 12, 13, 18, 20

weisingli 20
Eunotiaceae 11, 12, 18, 20
Eusphairica 28

distubula 28-30

F
Falcula 20
Forcepia 227, 228, 231, 238, 241, 242
Forcepia (Forcepia) 231, 242
acanthostylosa 228, 242
elvini 231, 242
forcipula 241
hartmani 235, 238, 242
hymena 238, 241—243
Japonica 231
macrostylosa 233
macrotylota 242
topsenti 235
Forcepia (Leptolabis) 231, 242
Frustulia 20

G
Gadomus 420-423
colletti 425
pepperi 417, 425, 426
sp. cf. colletti 417, 425, 426, 452
Glossochilus 152
Glossodoris 84
dollfusi 116, 119
Gorgoniidae 71
Gossypium spp. 249
Gotlandochiton 26
Gotlandochitonidae 24

H

Habracanthus 148
Haliporoides sibogae 412
Hansteinia 148
Haplomacrourus 472, 473

nudirostris 417, 426, 472
Hawthorneachiton lowenstami 24
Hemerobius conformis 289
Hemibagrus 125, 127, 129, 132, 138, 140

baramensis 127, 140

guttatus 127, 140

imbrifer 126, 127, 140

maydelli 128, 129

menoda 130, 132, 136, 138

microphthalmus 129, 133, 134

olyroides 127, 140

peguensis 132, 136-138

punctatus 129, 137-139

sabanus 127, 140

variegatus 127, 139, 140

wyckioides 129, 134
Hemidactylus

frenatus 403

garnoti 403
Heterocampa 20
Heteroxenia

capensis 93

uniserta 93
Hevea brasiliensis 249
Hinnites multirugosus 228
Hoplostethus atlanticus 413
Hygrophila auriculata 155
Hymenocephalus 426, 473, 478

aterrimus 417, 473-476

longibarbis 414, 474, 475

longiceps 475

nascens 417, 475

spp. 417
Hymenogadus (Hymenogadus) 473
Hyomacrurus 485
Hypoestes 146, 148, 152

aristata 144, 148, 155

forskaolii 144, 148, 155

triflora 156
Hypselodoris 116, 119, 123

babai 120, 122, 123

bullocki 122, 123

dollfusi 111, 116, 119

fucata 119

kaname 119

koumancensis 119

I

Ibacus 412

Idiolophorhynchus 500
andriashevi 500

Inuda 77,78, 82-84

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554 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 52

luarna 77, 78, 82, 84
Inudinae 82
Isididae 71
Tsoglossa 148
grandiflora 148
hypoestiflora 148
ovata 148
Italochrysa 251
Ivoechiton 28

J

Justicia 145, 148, 150, 152
adhatoda 148
anagalloides 156
betonica 144, 148, 150, 156
carnea 148
exigua 156
flava 150, 156
glabra 156
matammensis 156
odora 144, 150, 156
petiolaris 144, 150
sect. Harnieria 150
sect. Tyloglossa 150
trinervia 148

K

Keroeididae 71

Kindbladochiton 28

Kumba 486

Kuronezumia 476, 477, 486
bubonis 417, 476, 478
leonis 417, 477, 478
pallida 477
spp. 426

L
Lampophyton 66, 70, 87
planiceps 66, 68, 71
Lanthanocephalus 88
clandestinus 88, 89, 94
Leandra subulata \07
Lepidagathis 146
Lepidoptera 248
Lepidorhynchus 478
denticulatus 414, 417, 479
Leptogorgia 72
Leptolabis 227, 242
Leptophyton 87

Leucochrysa (Leucochrysa) 279, 302

ampla 249, 280, 282

boxi 249, 281, 284
bruneola 281, 283
pretiosa 249, 281-283
varia 249, 280, 282
walkerina 282, 283
spp. 248

Leucochrysa (Nodita) 250, 280, 284, 300, 302,

307
affinis 249, 287, 291, 293, 295, 299, 301
aleura 307
amazonica 284, 287
apicata 289
aurantiacus 249
barrei 249, 293, 295
camposi 249, 287, 301
clepsydra 249, 288, 295, 297
confusa 249, 296
cornuta 249, 288, 297
cruentata 249, 287-289, 295-297
forciformis 249, 297, 298
forcipata 297
furcata 249, 287, 297, 298, 304
gossei 289, 291, 296
guataparensis 249, 287, 291, 293, 295, 299
heriocles 249, 289
ictericus 249, 295, 300, 302, 308
lineata 303
incognita 249, 300
indiga 289
interata 249, 301
intermedia 249, 290, 292
internata 293
lancala 249, 291, 293, 299
lateralis 249, 291
lenora 297
lineata 249, 287, 292, 302, 304, 306, 308
maculata 249, 296
maculosa 302
marginalis 249, 290, 292, 294
marquezi 249, 293
melanocera 249, 287, 292, 293, 295
michelini 249, 287, 292, 302, 303, 308
morrisoni 307
nictheroyana 297
parallela 249, 293, 301, 304
postica 289, 302
retusa 249, 302, 304, 309
robusta 249, 305, 306
rodriguezi 249, 287, 292, 294, 300, 302
santini 249, 305, 306, 309
scomparini 249, 287, 306, 308
squamisetosa 249, 307

INDEX

tabacinus 249, 287, 292, 302-304, 308

tenuis 249, 308
trifurcata 307
vignisi 249, 309
virginiae 291
vittatus 249, 287
walkerina 249
Leucochrysini 279
Lissodendoryx 238
firma 238, 242
kyma 241
Lucigadus 479
denticulatus 408
microlepis 414, 480, 481
nigromaculatus 408, 481
Lycodon 397, 398, 403
aulicus 402-404
aulicus/capucinus 397, 398
fasciatus 397, 398, 403, 404
Jara 397, 398, 403, 404
kundui 397, 398
laoensis 397, 398, 403, 404
tiwarii 398, 404
travancoricus 398, 404
zawi 398, 402-404

M
Mackaya bella 156
Macrones 125
corsula 130, 136
menoda 130, 132, 136
microphthalmus 133
peguensis 136
punctatus 137
trachacanthus 130
Macrouridae 407, 426
Gadoidea 418
Macrourinae 426
Macrourcidae
Macrouroidea 418
Macrouroides inflaticeps 418
Macrouroididae
Macrouroidinae 426
Macrouronus
novaezealandia 418
spp. 418
Macrourus nigromaculatus 408
Macruronus novaezelandiae 479
Mahia matamua 449
Malacacanthus 93
capensis 71, 87, 89, 93, 94
rufus 93

Malacocephalini 485
Malacocephalus 480, 482
hawaiiensis 482
laevis 482, 483
luzonensis 482
nipponensis 482
Malus pumila 249
Mataeocephalus 483
acipenserinus 417, 484
sp. 417, 484
spp. 417
Mattheviidae 26
Species A 26
Megalochlamys 152
Melastomataceae 33, 514, 539
Melithaeidae 71
Merlucciidae 418
Mesobius 485
antipodum 417, 485
Metalcyonium 160, 198
Metarungia 144, 150
longistrobus 150
Miconia 33, 34, 39, 53
acanthocoryne 53
acuminifera 48
calocoma 34, 37
chiriquiensis 40
colliculosa 33, 34, 37
correae 37,39
crocata 39, 40
iteophylla 44
Jefensis 43, 44
ligulata 44
moril 44, 46, 48
peltata 44
rubens 40
rupticalyx 37
schnellii 50, 51
talamancensis 48, 50,51
vestita 51, 53
wagneri 37
zemurrayana 39
Mugil corsula 132
Mystus 125, 129, 138
corsula 130
gulio 138
krishnensis 128, 130
malabaricus 128
maydelli 128
menoda 130, 132, 136

menoda menoda 130, 136, 137, 138

menoda microphthalmus 133

555

556 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 52

menoda trachacanthus 130
microphthalmus 133
peguensis 136

punctatus 128, 130, 137
trachacanthus 130

sp. 128

N

Nacarina 251,255
aculeata 249, 253, 254
cordillera 255
gladius 249, 253, 254, 256
lavrasana 249, 253, 255
panchlora 249, 252, 253
pletorica 249, 252, 253
sagitta 249, 253, 255, 256
wagneri 253

Naviculaceae 11

Nelsonia canescens 156

Nephtheidae 71

Neula 279

Neuracanthus 152

Nezumia 476, 486
coheni 417, 487, 488
condylura 491
evides 49]
kapala 417, 488, 489
namatahi 417, 488, 489
propinqua 417, 490, 491
toi 489

Nidalia 160, 198
grayi 210

Nidaliudae 71

O

Odontomacrurus 491
murrayi 414, 492

Odontonema 150

Ophidocampa 20

Oplonia 150

Optonurus denticulatus 408

Oreosomatidae 413

P
Paleochiton
kindbladensis 24
siskiyouensis 24, 28
Paracetonurus sp. 435
Paralemnalia spp. 215
Paraminabea 71, 163, 202
Pennatulacea 209

Peristrophe 146, 148, 152
paniculata 156

Peronia 11, 18, 20
brasiliensis 12, 15, 18, 20

Phaulopsis imbricata 156

Phrynobatrachus 55, 56, 58-60, 62, 63

cricogaster 55
dendrobates 60, 64
irangi 56, 58-60, 62, 63
kinangopensis 60, 64
krefftii 56, 58-60, 63, 64
natalensis 60, 64
parkeri 60, 64
parvulus 60, 64
perpalmatus 60, 64
plicatus 60, 64
versicolor 60, 64
Phrynodon 55
Phyllidia varicosa 180
Phyllidiella pustulosa 180
Pieterfaurea 71, 87
Pimelodes
carcio 132
corsula 132
menoda 130, 132
telgagra 132
telagra menoda 132
Plesiochrysa 263,277,278
alytos 249, 277, 278

brasiliensis 249, 262, 277-279, 302

elongata 249, 277, 278
Plexauridae 71
Polyplacophora 24
Primnoidae 71
Pseuderanthemum 150, 152

hildebrandtii 150

laxiflorum 150

subviscosum 150
Pseudococcidae 248
Pseudonezumia 435
Psidium guajava 249

Quercus bumelioides 53

R

Razisea 148

Rhinacanthus 148, 150, 152
gracilis 144, 150, 152, 156
gracilis var. gracilis 152
gracilis var. latilabris 152

Rita 125

INDEX

Rostanga 1, 2,5,7
5

byga 1,3, 5-7
pulchra 2,6, 7,9
rubra 7

Rostangidae 2
Rostroconchia 28
Conocardioida 28
Rouxia 20
Ruellia
cordata 156
patula 156
Ruellieae 146
Ruelliopsis 152
Rungia 150
laeta 150
parviflora 150
pectinata 150
repens 150
Ruspolia 150, 152
hypocrateriformis 152
hypocrateriformis var. australis 152
seticalyx 152, 156
Ruttva 150, 152
fruticosa 152
ovata 152

S
Sarcophyton 72
Schoutedenella 55
Semiorbis 11
Simpsonella 72
Sinularia 72
Siphonoglossa 145, 148
linifolia 145
Sperata 125
Sphagemacrurus 492
decimalis 493
pumiliceps 493
richardi 417, 493
Sphenomorphus maculatus 403
Squalidae 413
Stenostephanus 148
Studeriotes 94
mirabilis 94
Syrphidae 248

T
Tessmannianthus carinatus 44
Thairoplax 24, 26
merriami 24, 26, 28
pelta 26
Thunbergia alata 156

Thysanoptera 248

S57

Topobea 97, 98, 100, 511, 512, 514, 534, 539,

547

aeruginosa 514, 516

albertieae 512,517,518

allenii 546, 547

amplifolia 518, 520

arboricola 98, 100, 102

brenesii 514, 520

caliginosa 100, 102

calophylla 521

calyeularis 322-523. 537.547;

cooperi 546, 547

cordata 102, 104

crassifolia 105, 107

dimorphophylla 523,525, 530, 544

dodsonorum 514, 526

durandiana 514, 533, 534

floribunda 539

fragrantissima 526, 527

gerardoana 527,529

hexandra 44, 102, 107, 108

inflata 542

intricata 525,529, 530

laevigata 523, 531

lentii 532

maurofernandeziana 514, 533, 534

mcephersonii 534, 535

membranacea 537, 539

multiflora 520, 523, 535, 537

parasitica 512, 537-539

parvifolia 539, 540

pitteri 540

pluvialis 541, 542

praecox 537,539

regeliana 537, 539

rhodantha 539

rosea 546, 547

rupicola 539

standleyi 542

suaveolens 543

subscaberula 520

superba 520

tetramera 512, 525, 543

urophylla 546, 547

watsonii 514, 532, 534, 547
Trachonurus 407, 494

gagates 417, 494, 495

sentipellis 417, 494, 495

villosus 495

yiwardaus 494

558 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 52

Trachyrincidae
Trachyrincinae 426, 500
Trachyrincus 407, 500
aphyodes 500
longirostris 417, 500
Tubastraea faulkneri 216
Tyrinna 84

V

Ventrifossa 480, 496
fasciata 481
fusca 497
Johnboborum 417, 496, 497
macropogon 499
misakia 497
nigrodorsalis 414, 497
nigromaculata 482
paxtoni 417, 499
saikaiensis 499

Ventrifossa (Sokodara) 497

Verconia 84

Verseveldtia 71, 87

Virgularia 72, 222

Ww
Wanwanella 28
Wanwanoidea 28
Wilsa 227, 228, 241
hymena 227, 228, 238, 241-243

X

Xenia uniserta 93
Xeniidae 71

Z
Zea mays 249

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